Identification of boreal forest stands with high herbaceous plant diversity using airborne laser scanning

Boreal forest stands with high herbaceous plant species diversity have been found to be one of the main habitats for many endangered species, but the locations and sizes of these herb-rich forest stands are not well known in many areas. Better identification of the stands could improve both their conservation and management. A new approach is proposed here for locating the mature herb-rich forest stands using airborne laser scanner (ALS) data and logistic regression, or the k-NN classifier. We show that ALS technology is capable of distinguishing the ecologically important herb-rich forests from those growing on less fertile site types, mainly on the basis of unique but quantifiable crown structure and vertical profile that characterise forests on high fertility sites. The study site, Koli National Park, is located on the border of the southern and middle boreal vegetation zones in Finland, and includes 63 herb-rich forest stands of varying sizes. The model and test data comprised 274 forest stands belonging to five forest site types varying from very fertile to poor. The best overall classification accuracy achieved with the k-NN method was 88.9%, the herb-rich forests being classified correctly in 65.0% of cases and the other forest site types in 95.7%. The best overall classification accuracy achieved with logistic regression was 85.6%, being 55.0% for the herb-rich forests and 94.3% for the other forest site types. Both methods demonstrated promising potential for separating herb-rich forests from other forest site types, although slightly better results were obtained with the non-parametric k-NN method, which was capable of utilising a higher number of explanatory variables. It is concluded that ALS-based data analysis techniques are applicable to the detection of mature boreal herb-rich forests in large-scale forest inventories.     

Habitat prediction model for three-toed woodpecker and its implications for the conservation of biologically valuable forests

We studied habitat selection of three-toed woodpecker (TTW) in relation to forest structural variables, habitat types and hydrological conditions in north-eastern Poland. Based on known locations of 34 breeding pairs and the data on location of forest areas with high conservation value (referred to as Biologically Important Forests), we created a habitat prediction model for TTW using a Maxent algorithm. We found that this species most often selected breeding sites characterized by high contribution of old-growth stands with uneven vertical structure and considerable amount of very old trees of previous generations, as well as boreal spruce-dominated bogs and riverine forests. Such sites were preferred regardless of their actual protection status, which leads to the following conclusion: although existing forest reserves are very important to three-toed woodpecker, suitable habitats with similar characteristics can still be found outside protected areas and are also utilized by the species. However, such sites outside reserves are threatened by fragmentation due to timber extraction. Therefore it is recommended to protect these sites by either including them into existing reserves or putting them under special management schemes.     

The effects of different types of management,functions, and characteristics of stands in Polish forests on the amount of coarse woody debris

In the modern forestry paradigm, many factors influence the amount of coarse woody debris (CWD). The present paper analyzes the effects of both local (national) programs (special functions of forests) and European programs (Natura 2000 sites), as well as the individual characteristics of forest stands. The study was conducted on 2,752 sampling plots distributed over an area of about 17,500 ha and located in lowland stands having a species composition typical of large areas in central Europe. Natura 2000 areas contained significantly more CWD (8.4 m³/ha) than areas not covered by the program (4.8 m³/ha). However, this is due to the fact that Natura 2000 sites involve well-preserved forest areas, such as nature reserves (26.6 m³/ha). In the managed forests that have been covered by the Natura 2000 program over the past several years, the volume of CWD has not increased. Forests with ecological and social functions differed slightly in the amount of CWD. More CWD occurred in protected animal areas (8.7 m³/ha) than in stands damaged by industry (3.9 m³/ha). Intermediate CWD levels were found in water-protection forests and in forests located around cities and military facilities. In managed forests, the lowest CWD volume was observed in middle-aged stands. The species composition of the stand had little effect on the volume of CWD. Only stands with a predominance of ash and alder had higher CWD levels (13.5 m³/ha). More CWD was found in stands whose species composition did not represent the potential site quality (6.4 m³/ha) than in habitats with the optimum species composition (3.8 m³/ha). CWD volume should be systematically increased taking into consideration local natural conditions. Such efforts should be focused on particularly valuable regions, and especially on Natura 2000 sites, where the threshold values reported from other European forests should be reached. Leaving some trees to die naturally and retaining reasonable amounts of such trees ought to be incorporated into CWD management practice in Poland.     

Human impacts drive forest structure and diversity. Insights from Mediterranean mountain forest dominated by <Emphasis Type="Italic">Abies pinsapo</Emphasis> (Boiss.)

Environmental factors and land-use are likely interacting to drive forest structure and species diversity, making it difficult to disentangle their separate impacts. Both components influence the resilience and stability of mountain forests in the Mediterranean Basin, since secular land-use has shaped stands structure and forests dynamics are constrained to the environment. Approaches covering environmental factors and contrasting land-uses are still needed to understand their implications on forest dynamics. We investigated patterns of environmental variables, stand structure and biodiversity in forest dominated by Abies pinsapo for 61 stands over their whole range. In addition, since land-use by local inhabitant could be a key factor, logging intensity was quantified by stumps basal area and recent land-use history was investigated. Partial redundancy analysis (pRDA) was conducted to assess to which degree the structural attributes are related to environmental characteristics and/or the land-use in the main A. pinsapo forests. Environmental characteristics accounted for 13% of the total variance, while the contrasting land-use (Spain vs. Morocco) and the stumps basal area, as indicator of human logging, accounted for 23% of the total variance. Trees species diversity was near 5 times higher in Moroccan stands than in stands from south Spain. According to recent human logging, Spanish stands have been strictly protected, but Moroccan stands showed widespread axe-logging by local inhabitant over the smaller tree sizes. Our results suggest that the preservation of a minor perturbation regimen by local populations enhance and help to preserve the whole biodiversity of this relict habitat.     

Relative importance of thinned and clear-cut stands for bryophyte diversity on stumps

The interest in harvesting logging residues in the form of stumps has recently increased markedly in the Nordic countries of Europe due to the steadily growing market for bio-fuel from forestry. Yet, stumps are currently harvested without a solid knowledge of their importance for forest organisms. To assess the relative importance of thinned, clear-cut, and burned clear-cut stands for the diversity of bryophytes on stumps we investigated bryophyte species composition and richness on 755 spruce stumps in 27 forest stands in southern-boreal Finland. Stumps were sampled within both wet and dry sites. For comparison we also surveyed bryophytes on 669 ground plots in the same stands to assess if patterns were consistent regardless of substrate type.Stand type (i.e. thinned, clear-cut, or burned clear-cut) was the main predictor of both species richness and composition on stumps. Stumps in thinned stands were more species rich than in clear-cuts, and the lowest richness was found on stumps in burned clear-cuts. Differences in species composition were explained by higher frequency of occurrence of species in thinned than in clear-cut stands. Thus, stumps in clear-cut stands tended to host a subset of species from the thinned stands, but there was also a pattern of turnover where different species were present in thinned and clear-cut stands, respectively. The results were similar for bryophytes on ground indicating that canopy cover strongly affected the diversity of bryophytes regardless of substrate.We conclude that stump harvest in open, clear-cut stands will probably have minor effects on bryophyte diversity on dead wood in the managed forest landscape. Conservation efforts in thinned stands (e.g. retention and creation of dead wood) may be important for promoting the diversity of bryophytes in managed forests.     

Retention of aspen (Populus tremulae) at final cuttings - The effect of dead wood characteristics on saproxylic beetles

To preserve biodiversity in managed forest landscapes dead and living trees are retained at final cuttings. In the present study we evaluated the effect of these practices for saproxylic (wood-dependent) beetles inhabiting dead aspen trees (Populus tremulae). For saproxylic beetles, tree retention at final cuttings can be expected to be especially valuable for species adapted to sun-exposed dead wood, a substrate that only rarely occurs in well managed forest stands. Therefore, the current evaluation was conducted as a comparison of species richness, species density (number of species per sample), assemblage composition and occurrence of individual species between clear-cuts, where aspen trees were retained, and closed forest stands with aspen trees. The study was conducted in central Sweden and the beetles were sampled by sieving of bark from CWD (coarse woody debris) of aspen. There was no significant difference in rarefied species richness between forest and clear-cut sites. Species composition differed significantly between the two stand types. Generalized linear mixed-effects models predicted the species density to be 34% lower in CWD objects in forest sites than on clear-cuts. This pattern could partly be explained by differences in CWD diameter, decay class and bark types between the two stand types (clear-cut/forest). Stand type was a significant predictor of occurrence in individual CWD objects for 30% of analysed individual beetle species. For all species except one, the variable stand type predicted higher occurrence on clear-cuts than in forest stands. To conclude, our results demonstrate that retention of aspen on clear-cuts contributes to population recruitment of a different assemblage of species than CWD within stands.     

Modelling coarse woody debris dynamics in even-aged Scots pine forests

Coarse woody debris (CWD) has become an important component in the study of forest ecosystems, being a key factor in the nutrient cycle as well as a habitat for many species. CWD dynamics varies greatly from primeval to managed forests. To assess the CWD dynamics, a chronosequence trial was established in two Scots pine forests in the Central mountain range in Spain. Although, the shelterwood system has been applied in both forests, one has received more intensive silviculture, whereas in the other, regeneration has been much more gradual and the thinning regime has not been so intensive. In order to inventory CWD, five decay classes and four categories according to size and CWD type (stumps and fallen logs or branches) were defined. The volume of branches and logs (estimated from length and mean diameter) and the number of stumps by size class and decay class are used to characterise the CWD. The most notable differences between the two silvicultural systems can be appreciated in the graph as peaks for temporary distribution of larger logs and stumps when intensive silviculture is applied. The CWD observed in the forests studied is mainly produced by logging. The maximum volume of logs and branches above 5 cm in diameter is 43.25 m³/ha after regeneration felling in the first forest, whereas in the other, a maximum of 16.30 m³/ha is reached at 60 years, just after thinning. Large stumps (diameter equal or greater than 30 cm) make up an important part of CWD biomass in these forests just after the regeneration felling. A model was developed to predict the changes in CWD quantity and quality distribution over time as well as predicting the effect of different silviculture options on CWD dynamics. The model integrates two different processes: the CWD inputs (which may be continuous or instant), and the decay process, modelled through a Richards–Chapman function. The average lifetime of CWD obtained ranges from 30 years for stumps over 30 cm in diameter to 8 years for logs with a diameter less than 10 cm.     

Regeneration of mixed conifer forests under group tree selection harvest management in western Bhutan Himalayas

Group selection tree harvest has been proposed as an ecologically sustainable silvicultural technique in mixed conifer forests of the western Bhutan Himalayas. To evaluate this silvicultural technique, we studied the ecological consequences of a group selection tree harvest in mixed conifer forests by assessing 127 circular plots (71 in logged and 56 in unlogged stands) in two forest management units (FMUs). Tree species composition and diversity were similar between logged and unlogged stands. Seedling density and height growth vary by species and were influenced by logging and microsites, with generally taller seedlings found in the logged versus unlogged stands. Early successional shade-intolerant species colonized logged stands. Seedlings growing on bare soil scarified by harvesting had medium vigour while seedlings growing on bryophyte mats showed good vigour in both logged and unlogged stands. Moist sites with a northerly aspect supported profuse conifer seedling regeneration, compared to sites with a dry southerly aspect. Damage to conifer seedlings from herbivore browsing was minimal. Conifer seedling density and height growth was negatively affected by competition from herbaceous vegetation, most notably Salvia officinalis. Group selection tree harvest in southern dry exposures in spruce-dominated stands is silviculturally unsuitable because it alters tree succession.     

Coarse woody debris and canopy cover in an old-growth Jeffrey pine-mixed conifer forest from the Sierra San Pedro Martir, Mexico

The cultural practices associated with Euro-American settlement in the United States have altered forest structure and ultimately changed fundamental ecosystem processes. Coarse woody debris (CWD) and canopy cover are recognized as having great importance for many wildlife species and ecological processes. Little information is available from forests on historical levels of canopy cover and CWD before European settlement. A great deal of uncertainty exists concerning the long-term role of fire and the dynamics of CWD, especially in forests that once experienced frequent, low-moderate intensity fire regimes. The objective of this study was to quantify CWD and forest canopy cover in an area where harvesting has never occurred and limited fire suppression began in the 1970s. This study was done in Jeffrey pine-mixed conifer forests in the Sierra San Pedro Martir (SSPM) in northwestern Mexico. Canopy cover, canopy closure, and CWD were sampled on a grid of plots. Average canopy cover was 26.8%, average canopy closure was 40.1%. A total of 102 CWD pieces were measured, and nearly half of the plots (45.7%) had no CWD present. Average CWD density, percent cover, volume, and weight were 108 pieces ha⁻¹, 1.5%, 47.5 m³ ha⁻¹, and 15.7 tonnes ha⁻¹, respectively. All of the CWD sampled were in the later stages of decay. Less than average values for CWD density, percent cover, volume, and weight were recorded in 57%, 64%, 67%, and 69% of the plots, respectively. CWD dynamics in forests that experience frequent, low-moderate intensity fires are fundamentally different than those having long-interval, high-severity fires. There was a large amount of variability in all CWD and forest canopy cover measurements taken from Jeffrey pine-mixed conifer forests in the SSPM. Spatial heterogeneity in forest structure should be included in the desired conditions of xeric, pine-dominated forests in the United States that once experienced frequent, low-moderate intensity fire regimes. It should be noted that heterogeneity by itself may not lead to sustainable forests unless that heterogeneity includes stand structures that are resistant/resilient to high-severity fire, drought, insects, and disease.     

Disturbance changes arbuscular mycorrhizal fungal phenology and soil glomalin concentrations but not fungal spore composition in montane rainforests in Veracruz and Chiapas,Mexico

Mexican montane rainforests and adjacent disturbed areas were studied for disturbance-related spatio-temporal changes to the arbuscular mycorrhizal fungal (AMF) community and soil glomalin concentration. The AMF community functions to both improve plant growth and soil conditions and is thus an important component to the restoration of this forest type to disturbed areas. The study areas included mature rainforests that were converted to pine forests, milpas, pastures and shrub/herbaceous plant communities via burning and logging. Seasonal patterns in AMF spore species richness and sporulation significantly differed across disturbance types at two of the three sites surveyed. Contrasting patterns of sporulation among AMF families across different disturbance types helped to explain how species richness and composition were maintained despite dramatic changes to the host plant community. Meaning, in most cases, disturbance induced changes in when different AMF taxa sporulated but not what taxa sporulated. Only conversion from mature pine–oak–Liquidambar–Persea forests to pine-dominated stands severely reduced AMF spore richness and total sporulation. Surprisingly, in pine-dominant stands no concomitant negative impacts on soil glomalin (MAb32B11 immunoreactive soil protein) concentrations were detected. However, soils of mature forests containing no pines had the highest concentration of glomalin. Conversion to pasture and milpa (diverse cornfield) had a strong negative impact on the concentration of soil glomalin concentrations. In sharp contrast, the same disturbance types improved AMF sporulation and AMF spore richness. It appears that disturbance type, and not AMF community measures used herein, best predicts changes in soil glomalin concentration.     

Response of fungal and plant communities to management-induced overstorey changes in montane forests of the Western Carpathians

The effect of forest management on biodiversity is a crucial issue for sustainable forestry and nature conservation. However, the ways in which management affects macrofungal and plant communities and diversity of mountain temperate forests still remain poorly understood. We performed a random sampling stratified by stand age and stand type on the sites of temperate montane fir–beech forests. Diversity of macrofungi and the vascular plant understorey in beech- and spruce-dominated managed stands was investigated and compared to primeval forests located in the Po?ana Biosphere Reserve, Western Carpathians. Both the vascular plant and the macrofungal communities were altered by management, and the response of the macrofungal species (especially wood-inhabiting fungi) was more pronounced in terms of species composition change. Species turnover evaluation seems to be an important tool of forest natural status assessment, because alpha diversity did not change as much as species composition. Certain species of Carpathian primeval forests were confirmed as good indicators for natural forest change; others were proposed. Species pool and mean number of species per plot were the highest in unmanaged fir–beech forests, and species diversity significantly decreased in spruce plantations. The number of species decreased significantly due to the change of canopy tree species composition only in the macrofungal communities. As an outcome for forest management, we recommend keeping mixed forests involving all natural tree species and providing at least a minimal amount of dead wood necessary for wood-inhabiting organisms and leaving some area of unmanaged natural forests within complexes of managed stands.     

Epiphytic lichen diversity in late-successional Pinus sylvestris forests along local and regional forest utilization gradients in eastern boreal Fennoscandia

Pinus sylvestris-dominated forests have been heavily utilized across all of boreal Fennoscandia and the remaining natural forests are generally highly fragmented. However, there are considerable local and regional differences in the intensity and duration of past forest utilization. We studied the impact of human forest use on the diversity of epiphytic and epixylic lichens in late-successional Pinus sylvestris-dominated forests by assessing species richness and composition along both local and regional gradients in forest utilization. The effects of local logging intensity were analysed by comparing three types of stands: (i) near-natural, (ii) selectively logged (in the early 20th century) and (iii) managed stands. The effects of regional differences in duration and intensity of past forest use were analysed by comparing stands in two contrasting regions (Häme and Kuhmo–Viena). The species richness of selectively logged stands was as high as that of near-natural stands and significantly higher in these two stand categories than in managed stands. Species richness increased with the density of small understorey Picea, which correlated strongly with decreasing intensity of local forest use and increasing structural complexity of selectively logged and near-natural stands. Stands in the Häme region hosted a lower number of species, and were less likely to host many old-growth indicator species than the Kuhmo–Viena region, suggesting that species have been lost from stands in the Häme region due to a longer history of intensive forest use. We conclude that selectively logged stands, along with near-natural stands, are valuable lichen habitats particularly for species confined to old-growth structures such as coarse trees and deadwood. In landscapes where natural forests have become fragmented, the management or restoration of the remaining late-successional Pinus-dominated forests, e.g. through the use of fire, should be carefully planned to avoid adverse effects on lichen species richness.     

Stocks and decay dynamics of above- and belowground coarse woody debris in managed Sitka spruce forests in Ireland

Coarse woody debris (CWD) has become recognised as an important component of the carbon (C) pool in forest ecosystems. In Ireland, managed Sitka spruce (Picea sitchensis (Bong) Carr.) forests account for 52.3% of the total forest estate. To determine the stock and decay dynamics of above and belowground CWD, field surveys using fixed area sample plots, were conducted in six even-aged Sitka spruce stands, representing the young, intermediate and mature stages of a typical commercial rotation. The volume, mass, density loss and C:N ratio of all CWD types (logs, stumps, and coarse roots) were determined using a five-decay class (DC) system. The decay rates and half life of CWD was also determined. To estimate CWD coarse root mass; roots associated with stumps classified in different decay classes were excavated. The coarse roots were categorised into small (2-10 mm), medium (10-50 mm) and large (>50 mm) diameter classes.CWD C-mass ranged from 6.98 to 18.62 Mg ha⁻¹ and was highest in an intermediate forest (D35), while the aboveground volume varied from 6.31 to 42.27 m³ ha⁻¹. Coarse roots accounted for 21% to 85% of the total CWD C-pool in the surveyed stands. The total CWD C-mass was poorly correlated with the number of thinning events (R² = 0.29), when data from D35 was excluded. The density loss was significant in logs (45%), stumps (58%), and small- (38%), medium- (50%) and large roots (38%) as decay progress from DC 0 to 4. There was a 46%, 41%, 51%, 72% and 57% decline in C:N ratio of logs, stumps, small-, medium- and large roots, respectively, as decay progressed from DC 0 to 4. The density decay rates were 0.059, 0.048 and 0.036 kg m⁻³ year⁻¹ for logs, stumps and coarse roots, respectively. The size classification of roots did not significantly affect their decay rate. The half life (50% decomposition) of CWD was estimated has 12-, 14- and 19 years for logs, stumps and roots of Sitka spruce. Regression curves showed a strong correlation between the density and C:N ratio (R² = 0.69, 0.74 and 0.93 for logs, stumps and coarse roots, respectively). The long term storage of C and its slow rate of decomposition make CWD a vital structural and functional component of the CWD C-pool and a major controller of forest ecosystem C-retention.     

Assemblages of wood-inhabiting fungi related to silvicultural management intensity in beech forests in southern Germany

Current silvicultural treatments in beech forests are aimed at achieving thick logs without discoloured hardwood. Therefore intensive thinning is applied already in younger stands with the objective of large-sized trunks at an age of 100 years. However, this approach bears the risk that dead wood structures and broken trees are completely removed from the forest. The impact of three different silvicultural management intensity levels on wood-inhabiting fungi over decades was investigated in a large beech forest (>10,000 ha) in southern Germany in 69 sampling plots: A Intensive Thinning and Logging with high-value trees, B Conservation-Oriented Logging with integration of special structures such as dead wood and broken trees and C Strict Forest Reserves with no logging for 30 years. The analysis of community showed marked differences in the fungus species composition of the three treatments, independent of stand age. The relative frequencies of species between treatments were statistically different. Indicator species for naturalness were more abundant at sites with low silvicultural management intensity. Fomes fomentarius, the most common fungus in virgin forests and strict forest reserves, is almost missing in forests with high-management intensity. The species richness seemed to be lower where intensive thinning was applied (P = 0.051). Species characteristic for coarse woody debris were associated to low management intensity, whereas species with a significant preference for stumps became more frequent with increasing management intensity. A total amount of dead wood higher than 60 m³/ha was found to enable significantly higher numbers of species indicators of naturalness (P = 0.013). In conclusion, when applying intensive silvicultural treatment, the role of dead wood needs to be actively considered in order to maintain the natural biocoenosis of beech forests.     

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.     

Effects of management on coarse woody debris volume and composition in boreal forests in northern Sweden

Forest management practices have led to a reduction in the volume and a change in the composition of coarse woody debris (CWD) in many forest types. This study compared CWD volume and composition in reserves and two types of managed forest in the central boreal zone of Sweden. Ten areas were surveyed, each containing clear-cut, mature managed and old-growth stands, to determine the volume of standing and lying CWD in terms of species composition, decay class and size class. Volumes of CWD on clear-cuts and in mature managed forests were high compared with previous studies. Old-growth forests (72.6 m³ ha^?1) contained a greater volume of CWD than mature managed forests (23.3 m³ ha^?1) and clear-cuts (13.6 m³ ha^?1). Differences were greatest for the larger size classes and intermediate decay stages. Despite stand ages being up to 144 years, CWD volume and composition in managed forests was more similar to clear-cuts than to old-growth forests.     

Stand specific occurrence of coarse woody debris in a managed boreal forest landscape in central Sweden

Forests long subjected to management for timber production contain only a fraction of the volumes of coarse woody debris (CWD) found in pristine forests. This is a threat to many organisms that depend on CWD. Forest management practices have been altered to achieve an increase in the amounts of CWD. Few studies have attempted to analyze the occurrence of CWD at the landscape level. We studied the occurrence of CWD in stands of different ages and management background in a boreal forest landscape in central Sweden. Volume of CWD in unmanaged stands (nature reserves and set-asides) was twice that in managed stands. The composition of CWD was influenced by stand age and management regime. Standing CWD was more common in unmanaged stands than in managed stands. Pine CWD was particularly prevalent in young forest stands (8–59 years of age). Bark-covered CWD was most common on deciduous and spruce wood and uncommon on pine. Bark area in young forest stands was almost 10 times lower than that in other managed stands. Using the age distribution of stands, we estimated the volume and bark area of CWD in the landscape. Recent clear-cuts harvested in accordance with new management guidelines contained more early decay CWD per ha than old managed stands. Young forests covered over half the landscape and had significantly lower volumes of spruce and deciduous CWD compared with other stands. The consequences of these results for biodiversity-oriented forest management are discussed.     

Tree regeneration in partially cut conifer–hardwood mixed forests in northern Japan: roles of establishment substrate and dwarf bamboo

We assessed the relationship of the regeneration (seedling and sapling) densities of seven representative tall-tree species to the past partial cutting and current stand structure. We also determined if differences in tree regeneration were associated to establishment substrates (coarse woody debris (CWD) and root throws) and understory inhibitor vegetation (the dwarf bamboo species: Sasa senanensis). The study was conducted in 17 conifer–hardwood mixed stands in a heavy snowfall region in Hokkaido, northern Japan. The results suggest that stand structure, rather than logging intensity, is the primary factor influencing regeneration densities. Total conifer basal area was positively correlated with the regeneration density of individual tree species, including two conifers and three hardwoods. These patterns differ from those observed in old-growth stands in the region. A negative correlation between total conifer basal area and dwarf bamboo coverage suggests that the presence of dense conifer canopies causes an increase in regeneration density of tall-tree species by preventing domination of dwarf bamboo. Picea glehnii, a species that depends for its establishment strongly on CWD, has lower seedling and sapling density in stands with higher logging intensity. This seems to be a result of the decrease in the volume of CWD with increasing logging intensity in these stands. We suggest that both reducing logging intensity and retaining overstory conifers should be considered to develop a sustainable silvicultural system in this region. Providing sufficient CWD and root throws may also be important to ensure natural regeneration of tree species that require these as an establishment substrate.     

Understorey vegetation in young naturally regenerated and planted birch (Betula spp.) stands on abandoned agricultural land

The abandonment of agricultural lands in Northern and Eastern Europe increases the area covered by first generation forests, which are either formed as an outcome of secondary succession or established as plantations. However, questions remain as to how these new stands develop and what kind of species they favour, which in turn has impacts on their ecological and economical value. Our aim was to compare understorey vascular plant and bryophyte vegetation characteristics between naturally regenerated and planted birch stands on abandoned agricultural sites in Estonia, focusing on the aspects of species richness and forest understorey recovery. Species richness and diversity of vascular plants were similar in both stand types but the number of forest vascular plant species was significantly higher in naturally regenerated stands. The bryophyte layer of naturally regenerated stands had a higher species richness, diversity, and number of forest bryophyte species. The higher number of forest vascular plant and bryophyte species in naturally regenerated stands can be explained by the longer undisturbed succession period. The recovery of the forest understorey was unaffected by former agricultural land use (crop field or grassland). The influence of soil properties on the recovery of the forest understorey was not detected, but the number of vascular plant species that grow in forests as well as in grasslands was negatively correlated with distance from forest. Overall, understorey vegetation of natural and planted birch stands did not reveal substantial differences. However, in the case of vigorous natural birch regeneration in the vicinity of forest land, unassisted reforestation should be favoured.