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.     

Responses of polypore fungi following disturbance-emulating harvesting treatments and deadwood creation in boreal Norway spruce dominated forests

ABSTRACT

The emulation of natural disturbances in harvesting has become a widely accepted approach to reach ecologically sustainable forest management. The purpose of this study was to examine the responses of polypore fungi (species richness and composition) on four different cutting methods (selection, gap, patch, and clear-cutting) plus uncut controls, and the creation of high stumps and cut logs in combination with harvesting. Polypores were inventoried one year before, and two and seven years after cutting. A total of 71 polypore species were found, but no red-listed species were recorded seven years after cutting. Irrespective of the treatment, the variation in polypore richness was best explained by the number of deadwood objects in the stands. Species richness per deadwood object was the highest on natural logs. Species richness per volume of spruce deadwood was the highest on small-diameter deadwood and stumps, intermediate on natural logs and lower on cut logs and high stumps. These results indicate that cut logs created in harvesting cannot compensate the loss of deadwood in cutting and energy wood harvesting. Our results demonstrate the difficulty in rapidly creating habitats for red-listed polypores in commercially managed forests.     

A lack of large-diameter logs and snags characterises dead wood patterns in Irish forests

Dead wood is an important component of forest ecosystems and volumes vary depending on forest age, management intensity and productivity. This is the first large-scale study to quantify dead wood in Irish forests and to compare them to forests in other locations. We measured the volume and size distribution of logs, the density and size distribution of snags and the volume of dead wood contained in stumps in Oak (Quercus spp.) and Ash (Fraxinus excelsior) forests and in Sitka spruce (Picea sitchensis) plantations throughout Ireland. We also assigned each log, snag and stump to one of three decay classes (intact, part-rotted and well-rotted). We found no significant difference in log volume between any of the forest types. The majority (>90%) of logs were less than 20 cm in diameter, and large logs (>40 cm diameter) were scarce. We found a relatively high density of snags in all forest types but, as in the case of logs, over 90% of snags were <20 cm DBH and large snags (>40 cm DBH) were rare. The volume of dead wood contained in stumps was significantly higher in plantations than in Oak or Ash forests as a result of thinning and harvesting. Most logs and snags were moderately decayed but, in plantations, most stumps were intact. Log volume and the size of logs and snags were considerably lower than in old-growth forests in other regions. These patterns may reflect historical use of Irish forests for coppice and timber production. Management for biodiversity should aim to accelerate dead wood accumulation to increase the frequency of large-diameter logs and snags. Although management seeking to replicate the dead wood volumes of old-growth forests is ideal, it may be unrealistic in the short term.     

Immediate Effects of Logging,Mounding, and Removal of Logging Residues on Epixylic Species in Managed Boreal Norway Spruce Stands in Southern Finland

The immediate effects of bioenergy harvesting methods on epixylic species were studied in mature managed Norway spruce dominated forests in southern Finland. The treatments included logging, residue harvesting, and soil preparation as either mounding or mounding combined with stump harvesting. Altogether, 110 logs and 440 species sample plots on logs were inventoried before and after logging, and after soil preparation treatments. Logging decreased the cover and species richness in all epixylic species groups. The soil preparation decreased the cover of macrolichens, while stump harvesting was significantly most devastating both for cover and richness in bryophytes. We suggest that bioenergy harvesting critically affects the epixylic species, and therefore the use of less devastating methods and compensation for the loss of coarse woody debris is recommended on a landscape level.     

Drivers of bryophyte diversity allow implications for forest management with a focus on climate change

Understanding the major drivers of diversity in forests is crucial for ecologists, conservationists, and particularly forest managers. In most forested habitats, bryophytes are diverse and important primary producers. Here we report the first study of the relative importance of regional macroclimate as compared to local variables on abundance, species richness, and community composition of bryophytes growing in soil and in dead wood in a mountain temperate forest. Using PCA axes, we built predictor sets for macroclimate, microclimate, soil attributes, and dead wood availability. The explanatory power of each set was tested using variance partitioning. Abundance and species richness of soil bryophytes was best explained by microclimate, whereas the community composition did not distinctively differ between the environmental sets. In contrast, dead wood bryophyte abundance, species richness, and community composition was clearly driven by macroclimate. Among the single axes, the component represented best by soil moisture was the main driver for soil bryophyte abundance. In contrast, dead wood bryophyte abundance was mostly affected by components correlated with minimum global radiation and minimum temperatures as well as by the component represented by dead wood. The component represented by canopy openness was superior in explaining the community composition of both soil and dead wood bryophytes. We conclude that (1) dead wood amounts should be increased in closed stands to act as a buffer during climate changes, and (2) open canopy, which provides important habitats for soil-inhabiting bryophytes, should be provided by management with slow reforestation after natural or logging openings.     

Targets for maintenance of dead wood for biodiversity conservation based on extinction thresholds

Abstract

Forestry decreases the amount of dead wood, thereby threatening the persistence of many saproxylic (wood-living) organisms. This article discusses how targets for efforts to maintain and restore dead wood in managed forest landscapes should be defined. Several studies suggest extinction thresholds for saproxylic organisms. However, because the thresholds differ among species, the relationship between species richness and habitat amount at the local scale is probably described by a smoothly increasing curve without any distinct threshold. The most demanding species require amounts of dead wood that are virtually impossible to reach in managed forests. This means that unmanaged protected forests are needed. In managed forests, conservation efforts should focus on the landscape scale and on certain types of dead wood, but it is impossible to come up with any particular amount of dead wood that is desirable at the forest stand level.     

Legacies from natural forest dynamics: Different effects of forest management on wood-inhabiting fungi in pine and spruce forests

The species composition of wood-inhabiting fungi (polypores and corticoids) was investigated on 1138 spruce logs and 992 pine logs in 90 managed and 34 natural or near-natural spruce and pine forests in SE Norway.Altogether, the study included 290 species of wood-inhabiting fungi. Comparisons of logs with similar properties (standardized tree species, decay class, dimension class) in natural and managed forests showed a significant reduction in species number per log in managed spruce forests, but not in managed pine forests. The species number per log in managed spruce forests was 10-55% lower than on logs from natural spruce forests. The reduction was strongest on logs of large dimensions. A comparison of 200-400 spruce logs from natural and managed forests showed a 25% reduction in species richness corresponding to a conservative loss of ca. 40 species on a regional scale.A closer inspection revealed that species confined to medium and very decayed spruce logs were disfavored in managed forests, whereas species on early decay classes and decay generalists were unaffected. Similarly, species preferring large spruce logs were disfavored in managed forests. Forest management had strongest impact on low-frequent species in the spruce forests (more than 50% reduction), whereas common species were modestly affected. Corticoid fungi were more adversely affected than polypore fungi.These results indicate that wood-decaying fungi in pine forests are more adapted to forest disturbances than spruce-associated species. Management measures securing a continuous supply of dead wood are more important in spruce forests than in pine forests.     

Saproxylic beetle assemblages on low stumps,high stumps and logs: Implications for environmental effects of stump harvesting

The aim of this study was to evaluate how future harvesting of spruce low stumps as a renewable energy source, might influence the saproxylic (wood living) beetle fauna. Species richness, abundance and assemblages of saproxylic beetles were compared on low stumps, high stumps and logs of spruce on 10 clear-cuts in northern Sweden. The sampling was conducted with emergence traps and a total of 929 individuals and 120 species were collected. No significant differences in beetle abundance or species richness were detected between substrate types. However, there were clear differences in assemblage composition between all substrate types. Our results suggest that low stumps created at final felling support as many species and individuals per volume dead wood as other dead wood substrates and that they are therefore overlooked as substrates for wood living organisms. Consequently, harvesting of low stumps might have negative effects on the diversity of saproxylic beetles, as up to 80% of the dead wood remaining on clear-cuts is in the form of low stumps. Future studies should evaluate if compensatory measures can be performed to minimize negative impacts.     

A continuum of dead wood with various habitat elements maintains the diversity of wood-inhabiting fungi in an old-growth boreal forest

Evidence on habitat associations of threatened wood-inhabiting species in boreal forests may contribute to a better understanding of their ecology and conservation needs. We examined the diversity of wood-inhabiting fungal communities in an old-growth boreal forest with high substrate availability and continuity based on repeated surveys of fruit bodies. The number of species in morphological and functional groups in relation to coarse woody debris (CWD) attributes was estimated with generalized linear models. Additionally, we calculated species interaction networks of CWD attributes and fungal species. The composition of fungal communities was analysed using a non-metric multidimensional scaling with subsequent environmental fitting. Old conifer (especially spruce) logs and large aspen logs with branches represented the most important substrates for the red-listed species and the indicator species of old-growth boreal forests. Among “dynamic” CWD attributes such as time since tree death, decay class and stage of epixylic succession, the latter was the most important indicator of diversity of all species and their morphological and functional groups. The interaction network provided evidence of the importance of tree species diversity for fungal diversity. The composition of fungal communities was tree species specific and related to dynamic attributes, bark cover and diameter of logs. Our results suggest the importance of a continuum of dead wood from different tree species with a variety of niches such as branches, exposed wood, fragmented and complete cover of bark and patches of epixylic vegetation to maintain the assemblages of wood-inhabiting fungi in an old-growth boreal forest.     

海南霸王岭陆均松类热带山地雨林伐后林结构与物种多样性研究

霸王岭林区是海南岛最典型的热带山地雨林分布区.在该区,现存分布面积最广、资源数量最大的是热带山地雨林的各种伐后林.以霸王岭分布最广的以陆均松(Dacridium pierrii)为主的山地雨林的伐后林为对象,研究了其组成、径级、高度结构和物种多样性特征,对比分析了伐后林与原始林以及伐后林不同类型之间的异同.结果表明:原始热带山地雨林经采伐后中小乔木和灌木的优势度增大,而大乔木树种的优势度则明显下降.阳性树种的种类、数量和相对重要性随采伐破坏的程度而增加.伐后林中矮、小树木在林分中比例随采伐破坏的程度增加,而高、大树木,特别是进入主林层树木的比例则随采伐破坏的程度明显下降.从林分测树因子的变化来看,采育择伐和径级择伐的林分要优于大强度择伐和皆伐的林分.原始林的物种丰富度大于伐后林,但其多样性指数和均匀度指数大都小于伐后林.组成上与原始林相似性较大的伐后林是各种择伐后的林分,而皆伐后的林分与原始林在组成上已有很大差别.     

Bryophytes and decaying wood in Hepatica site-type boreo-nemoral Pinus sylvestris forests in Southern Estonia

The aim of the study was to establish the amount of decaying wood (logs and stumps) in various groups of Hepatica site-type pine forests of different age and management intensity and to analyse the composition of bryophytes in dependence of these factors. The average volume of CWD in old unmanaged forests was 47.5 m³/ha, which is rather well comparable with respective estimations from Fennoscandia. Reduced human impact contributes positively to the amount of CWD. Diversity of log diameter classes and decay stages is larger in old forests. Altogether 73 bryophyte species were recorded, 65 species on logs and 55 on stumps. Species richness on stumps was higher in managed forests than in unmanaged ones. At the same time, the species having high indicator value for man-cut stumps are very common species in boreal forests and grow on other substrata as well. Species composition and ecological conditions differed between stumps and logs. Logs are more humid microhabitats than stumps, therefore the occurrence of hepatics is more frequent on them. According to species composition on decaying wood the old unmanaged forests distinguished from others. As the differences of substratum characteristics were notable between old and young forests, the stand age described a considerable part of species variance on logs.     

Red-listed wood-inhabiting fungi in natural and managed forest landscapes adjacent to the timberline in central Sweden

Many wood-inhabiting fungi are today threatened as modern forestry practices drastically reduce the amount of dead wood available in various forest ecosystems. We investigated whether the occurrence of red-listed wood-inhabiting fungi differed between natural and managed forest landscapes adjacent to the timberline in the middle part of Sweden. We assessed whether environmental variables such as the degree of human impact, length of forest roads, dead wood volume and quality affected species richness and abundance. The effects of forestry on wood-inhabiting fungi have been assessed in several studies in lowland Swedish forests. Few studies have, however, been conducted in forest landscapes adjacent to the timberline in Sweden. This is potentially important since forests close to the Swedish mountains have been pointed out as one of few intact forest landscapes in Fennoscandia and they are subjected to increasing logging pressure. Similar to other studies, species numbers and abundances were positively correlated with larger volumes of logs in various decay stages. However, never shown previously, the length of forest roads was negatively correlated with species abundance and occurrence of red-listed species. We suggest that a low amount of forest roads can be used as a conservation indicator to localize still-intact forest landscapes.     

Effects on the flora in Norway spruce forests following clearcutting and shelterwood cutting

Clearcutting and shelterwood cutting of mature Norway spruce peatland forests were compared regarding the effects on the forest flora. Repeated observations of the field and bottom layer vegetation were made before and until 7 or 8 years after harvesting at four sites located along a gradient from southern to northern Sweden. Clearcutting resulted in a greater change of species composition compared with shelterwood cutting. Diversity, measured as Simpson's index, and species number per subplot were lower in the clearcut than in the shelterwood after 7 or 8 years. By applying Ellenberg's indicator values, it was concluded that shelterwood regimes may preserve species preferring shaded and moist conditions, whereas those species decreased after clearcutting. Species preferring high levels of nitrogen increased in the clearcut.According to both multivariate and univariate analyses, the vegetation changed in a similar direction at all sites, although the level of response differed considerably. Despite these similarities, there was a marked site effect, which was not surprising considering the large geographic variation between sites.It is concluded that shelterwood cutting might be a better alternative than clearcutting for forests on fertile peatland sites, with respect to conservation of vascular plants and bryophytes.     

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.     

Short term effects of alternative silvicultural treatments on stand attributes in irregular balsam fir-black spruce stands

The eastern Canadian boreal forest exhibits a specific disturbance regime where forest fires are less frequent than in the western part. This particularity may explain the abundance of irregular stands with distinct ecological features. To ensure sustainable forest management, these characteristics require the implementation of an adapted silviculture regime. In this context, two selection cutting methods were developed and compared with more conventional techniques, initially designed for cutting more regular stands of the boreal forest (cutting leaving small merchantable stems, careful logging preserving advance regeneration). The comparison focused on the capacity of treatments to maintain the primary attributes of irregular boreal forests, including complex vertical structure, abundant tree cover, species composition, and an abundance of dead wood. Mortality and regeneration processes were also compared.     

Epiphytic lichen diversity in old-growth and managed Picea abies stands in Alpine spruce forests

In the last decades, a large body of literature has grown to evaluate the impact of forest management on epiphytic lichens in boreal coniferous forests. However, information is still lacking on coniferous forests of the Alps. This study compares lichen diversity between spruce forest stands of four successional stages: (1) young, (2) intermediate, (3) mature forests managed for timber production with a rotation cycle of 120–180 years, and (4) old-growth protected forests. The emphasis was placed on the occurrence of nationally rare and calicioid species (lichens and fungi traditionally referred to as Caliciales, known to be indicative of forest age and continuity). For each forest successional stage, four plots were selected. In each plot, 7 spruce individuals were surveyed for epiphytic lichens according to a standardised sampling method. Species richness increased from young to mature stands, while no difference was detected between mature and old-growth stands. This pattern was also confirmed for rare and calicioid species which are, however, more frequent in old-growth stands. Differences in species composition were also found between the different forest successional stages. Mature and old-growth plots slightly overlap, indicating that to some extent comparable lichen assemblages could be found in these stands. A nested pattern of species assemblages was found, old-growth stands hosting most of the species which were also found in stands belonging to the previous forest successional stages. Our results support the hypothesis that the management regime applied to spruce forests of the Italian Alps renders mature stands managed for timber production somewhat similar to old-growth stands as lichen habitat. However, we found a higher complexity in old-growth forests, and many species of conservation concern clearly preferred old-growth stands. In this perspective, a further prolongation of the normal cycle it is likely to be a most favourable conservation-oriented management to be recommended at least within protected areas and Natura 2000 sites, where conservation purposes should receive a high priority.     

Foraging behavior of pileated woodpeckers in partial cut and uncut bottomland hardwood forest

In bottomland hardwood forests, partial cutting techniques are increasingly advocated and used to create habitat for priority wildlife like Louisiana black bear (Ursus americanus luteolus), white-tailed deer (Odocoileus virginianus), and Neotropical migrants. Although partial cutting may be beneficial to some species, those that use dead wood may be negatively affected since large diameter and poor quality trees (deformed, moribund, or dead) are rare, but normally targeted for removal. On the other hand, partial cutting can create dead wood if logging slash is left on-site. We studied foraging behavior of pileated woodpeckers (Dryocopus pileatus) in one- and two-year-old partial cuts designed to benefit priority species and in uncut forest during winter, spring, and summer of 2006 and 2007 in Louisiana. Males and females did not differ in their use of tree species, dbh class, decay class, foraging height, use of foraging tactics or substrate types; however, males foraged on larger substrates than females. In both partial cut and uncut forest, standing live trees were most frequently used (83% compared to 14% for standing dead trees and 3% for coarse woody debris); however, dead trees were selected (i.e. used out of proportion to availability). Overcup oak (Quercus lyrata) and bitter pecan (Carya aquatica) were also selected and sugarberry (Celtis laevigata) avoided. Pileated woodpeckers selected trees ≥50 cm dbh and avoided trees in smaller dbh classes (10–20 cm). Density of selected foraging substrates was the same in partial cut and uncut forest. Of the foraging substrates, woodpeckers spent 54% of foraging time on live branches and boles, 37% on dead branches and boles, and 9% on vines. Of the foraging tactics, the highest proportion of foraging time was spent excavating (58%), followed by pecking (14%), gleaning (14%), scaling (7%), berry-eating (4%), and probing (3%). Woodpecker use of foraging tactics and substrates, and foraging height and substrate diameter did not differ between recent partial cut and uncut forest. Partial cutting designed to improve or maintain habitat for priority wildlife did not affect pileated woodpecker foraging behavior or availability of selected trees compared to uncut forest in the short term.     

Retention patches maintain diversity of epiphytic and epixylic indicator lichens more effectively than solitary trees

The effectiveness of retention trees and patches in preserving diversity of nine epiphytic and epixylic old-growth forest lichens was studied in north boreal spruce forests in Finland. We compared (1) 7–8-year-old retention cuts, with at least 5–10 living or dead retention trees per hectare, (2) 10–12-year-old clear-cuts, with some scattered living and dead retention trees on the sites, (3) old-growth spruce forests, and (4) 7-8-year-old retention patches (0.06–0.45?ha) representing the original tree species composition of old-growth forests. The occurrence of indicator lichens was studied on 150 deciduous trees and snags in each forest category. The species richness was significantly higher in old-growth forests than in the clear-cuts and retention cuts, but did not differ between old-growth forests and retention patches. Only three species were found in clear-cuts and two in retention cuts. Foliose cyanolichens Leptogium saturninum and Nephroma bellum thrived on solitary retention trees, whereas humidity-requiring pin lichens from the genus Chaenotheca were found only in old-growth forests and retention patches. Our results suggest that the ability of epiphytic and epixylic species to survive on retained trees depends on several factors: (1) substrate quality (tree species, tree type and diameter of a tree), (2) environmental factors (e.g. humidity, slope exposition), and (3) morphological and physiological characteristics of species. Besides of substrate trees, the retained conifers (esp. spruce) seem to be important in retention patches to provide the shading necessary to maintain humidity.