Saproxylic beetles in high stumps and residual downed wood on clear-cuts and in forest edges

Abstract

This study evaluates how the placement and the different possible outcomes of a spruce retention tree affected species richness and assemblages of spruce-associated saproxylic beetles. In a field experiment in the boreal zone of central Norway, high stumps were created and compared with residual wood pieces (i.e. top boles with branches), in clear-cuts versus in forest edges. Flight interception traps were mounted close to the substrate. The results were analysed using rarefaction techniques, ordination (DCA) and anova. It was found that the placement of retention trees of spruce does matter: beetle assemblages were significantly different in stumps in the four treatments. For all species pooled, the species richness was higher in stumps in clear-cuts than in stumps or boles in the forest edge. Four red-listed species were more abundant near stumps than boles, and two red-listed species were more abundant in clear-cuts than forest edge. To cater for the variety of habitat preferences among forest beetles in managed forest, managers should leave both standing spruce trees (many of which will end up as windthrown, downed boles) and some high stumps (to secure some upright dead wood), in both exposed clear-cut and semi-shaded forest edge.     

Decaying wood and tree regeneration in the Acadian Forest of Maine,USA

We examined the effect of management history on the availability of decayed downed wood and the use of downed wood as a regeneration substrate in mixed-species stands in the Acadian Forest of Maine. Regeneration of red spruce (Picea rubens Sarg.), eastern hemlock (Tsuga canadensis (L.) Carr.), balsam fir (Abies balsamea L. Mill), and red maple (Acer rubrum L.) was quantified. Treatments included variants of selection cutting, commercial clearcutting (unregulated harvesting), and no harvesting for >50 years (reference). Area of wood substrate (wood ≥ Decay Class III and ≥10 cm on at least one end) was less in the commercial clearcut than in the reference; other treatments were not differentiated. Spruce and hemlock seedlings were found at higher densities on wood than paired forest floor plots of equal area, regardless of treatment. Conversely, fir and maple were less abundant on wood than forest floor plots in reference and selection treatments, but more or equally abundant on wood than forest floor plots in the commercial clearcut. These findings suggest that silvicultural treatment affects both the availability of decayed downed wood and seedling-substrate relationships, and that forest management in the Acadian Region should consider availability of downed woody material.     

Habitat associations of saproxylic beetles in the southeastern United States: A comparison of forest types,tree species and wood postures

Saproxylic beetles are highly sensitive to forest management practices that reduce the abundance and variety of dead wood. However, this diverse fauna continues to receive little attention in the southeastern United States even though this region supports some of the most diverse, productive and intensively managed forests in North America. In this replicated three-way factorial experiment, we investigated the habitat associations of saproxylic beetles on the coastal plain of South Carolina. The factors of interest were forest type (upland pine-dominated vs. bottomland hardwood), tree species (Quercus nigra L., Pinus taeda L. and Liquidambar styraciflua L.) and wood posture (standing and downed dead wood, i.e., snags and logs). Wood samples were taken at four positions along each log and snag (lower bole, middle bole, upper bole and crown) ∼11 months after the trees were killed and placed in rearing bags to collect emerging beetles. Overall, 33,457 specimens from 52 families and ≥250 species emerged. Based on an analysis of covariance, with surface area and bark coverage as covariates, saproxylic beetle species richness differed significantly between forest types as well as between wood postures. There were no significant interactions. Species richness was significantly higher in the upland pine-dominated stand than the bottomland hardwood forest, possibly due to higher light exposure and temperature in upland forests. Although L. styraciflua yielded more beetle species (152) than either Q. nigra (122) or P. taeda (125), there were no significant differences in species richness among tree species. There were also no relationships evident between relative tree abundance and observed or expected beetle species richness. Significantly more beetle species emerged from logs than from snags. However snags had a distinct fauna including several potential canopy specialists. Our results suggest that conservation practices that retain or create entire snags as opposed to high stumps or logs alone will most greatly benefit saproxylic beetles in southeastern 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.     

Development and activity of early saproxylic fungal communities in harvested and unmanaged boreal mixedwood stands

Limited scientific information is currently available regarding saproxylic fungal communities in the boreal forest of North America. We aimed to characterize the community development, richness and activity of saproxylic fungi on fresh wood in harvested and unmanaged boreal mixedwood stands of northwestern Québec (Canada). Fresh wood blocks (n = 480) of balsam fir (Abies balsamea (L.) Mill.) and trembling aspen (Populus tremuloides Michx.) were placed on the forest floor in a range of stand conditions (n = 24). Blocks were harvested every 6 months for up to 30 months and characterized for species composition and richness (PCR–DGGE, DNA sequencing), respiration, wood density and lignin and cellulose content. Colonization by a wide range of functional groups proceeded rapidly under different stand conditions. We detected a total of 35 different fungal operational taxonomic units, with the highest species richness at the wood block level being observed within the first 12 months. No differences in community composition were found between wood host species or among stand conditions. However, the variability in fungal communities among blocks (β diversity) was lower on trembling aspen wood compared with balsam fir and decreased over time on trembling aspen wood. Also, fungal activity (respiration and wood decomposition) increased on trembling aspen wood blocks and species richness decreased on balsam fir wood over time in partial-cut sites. The overlap in tree composition among stands, the high volume of logs and the recent management history of these stands may have contributed to the similarity of the saproxylic fungal community among stand types and disturbances.     

The choice of definition has a large effect on reported quantities of dead wood in boreal forest

A survey was conducted to assess the impact of the choice of definition on reported quantities of dead wood in Swedish forests, which to more than 90% are located in the boreal zone. The data collection was made on a subsample of the permanent plots of the Swedish national forest inventory. The objects included were standing dead trees and snags down to 5-cm diameter at breast height, dead lying stems and branches down to a threshold diameter of 1 cm and stumps down to a threshold diameter of 5-cm at normal stump height. Standing trees, snags and stumps were inventoried on 10-m radius circular plots while the downed objects were inventoried using both circular plots and line intersect sampling; thin objects (diameter 1–5 cm) were assessed only through line intersect sampling. The results showed that the estimated volume of dead wood was as high as 25 m³ ha^?1 when all components were included. With the standard Swedish definition, the corresponding estimate was only 10.9 m³ ha^?1, or 43% of the total value. Since definitions of dead wood vary greatly between countries we conclude that great caution must be exercised when figures are compared in connection with international reporting. For example, adding stumps to the Swedish definition would increase the amounts of dead wood from 10.9 to 15.7 m³ ha^?1, i.e. with 44%.     

Recovery of boreal forest structures near abandoned villages in Western White Sea Karelia,Russia

Regeneration of natural forest structures and dead wood has a key role in the long-term maintenance of biodiversity in most parts of the boreal zone. In order to obtain benchmarks for practical dead wood management rapidly, we utilised unintended historical experiments in land use by studying the recovery of forests surrounding abandoned villages along 2.5-km transects in NW Russia. The signs of past human forest use declined as a function of distance from villages. We found that the closest forests were 130 yr old or younger and contained very few elements typical of natural forests. Forests at a distance of between 1 and 1.5 km were older than 150 yr but showed marks of selective cuttings and had a less complete continuum of dead wood than forests that were 2–2.5 km from the villages. The most remote stands, which were generally older than 200 yr, predominantly had no signs of past forest use and had the most diverse dead wood composition. Our study shows that full recovery of the dead wood continuum in middle boreal forests is a slow process and also that less intensive past forest use can have a significant effect on the dead wood continuum.     

Distribution of saproxylic beetles in a recently burnt landscape of the northern boreal forest of Québec

This study evaluated the importance of burned habitat characteristics as well as the likely dispersal from specific habitats in the distribution of saproxylic beetles the same year as a fire occurred, in burned black spruce stands (Picea mariana [Mill] B.S.P.) in the northern boreal forest of Québec. The distribution of early post-fire saproxylic species was mainly driven by burned habitat attributes at the plot scale (0.04 ha), especially fire severity, suggesting that the effect of environment attributes can act at a relatively fine scale. Some xylophagous and most predaceous species were more abundant in severely burned stands whereas fire severity had the opposite effect on several common mycophagous species. The amount of newly fire-killed trees that could be used as breeding substrates in the burned stands had only a weak positive influence on these functional groups. The great majority of early saproxylic species were weakly associated with the distance from unburned forests or other recently burned patches that could act as potential “source habitats”. Indeed, these variables were of lesser importance than the attributes of the burned habitat. Woody debris that were already present in plots before the fire, potentially serving as local of source-populations for early colonizers, had virtually no influence on the local abundance of species. Many saproxylic species, including some true pyrophilous, clearly showed higher abundance as distance from unburned stands increased. This unexpected relation may reflect that dispersal of insects toward the burnt landscape very shortly after fire could be driven by the higher amount of volatiles released by severely burned forests, which are more likely as distance from unburned forest increased.     

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.     

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.     

Dead wood in European beech (Fagus sylvatica) forest reserves

Data were analysed on the volume of dead wood in 86 beech forest reserves, covering most of the range of European beech forests. The mean volume was 130 m³/ha and the variation among reserves was high, ranging from almost nil to 550 m³/ha. The volume depended significantly on forest type, age since reserve establishment and volume of living wood. More dead wood was found in montane (rather than lowland/submontane) reserves, longer-established reserves (time since designation) and reserves with higher volumes of living wood.

On average, fallen dead wood contributed more to the total dead wood volume than standing dead wood. The percentage of dead wood that was standing was almost twice as high in montane than in lowland/submontane forest reserves (45% versus 25%). The volume of dead wood at selected sites changed considerably over time. The fluctuations were significantly higher in lowland/submontane than montane reserves, possibly connected with differences in the disturbance regimes and especially damage caused by windstorms. In NW Europe, the blow down of formerly managed, even-aged stands led to extraordinary high volumes of dead wood shortly after reserve establishment.

The implications for forest management and biodiversity conservation are discussed. An increase in dead wood volumes must be carried out in accordance with the local/regional forest type and disturbance regime. Thus, in order to fulfil the requirements of as many wood-depending organisms as possible, it is important to preserve not only larger amounts of dead wood, but also dead wood of different types and dimensions as well as securing a long-term continuity of dead wood.     

Impact of silviculture on dead wood and on the distribution and frequency of tree microhabitats in montane beech-fir forests of the Pyrenees

In forest ecosystems, the level of biodiversity is strongly linked to dead wood and tree microhabitats. To evaluate the influence of current forest management on the availability of dead wood and on the abundance and distribution of microhabitats, we studied the volume and diversity of dead wood objects and the distribution and frequency of cavities, dendrothelms, cracks, bark losses and sporophores of saproxylic fungi in montane beech-fir stands. We compared stands unmanaged for 50 or 100 years with continuously managed stands. A total of 1,204 live trees and 460 dead wood objects were observed. Total dead wood volume, snag volume and microhabitat diversity were lower in the managed stands, but the total number of microhabitats per ha was not significantly different between managed and unmanaged stands. Cavities were always the most frequent microhabitat and cracks the least frequent. Dendrothelm and bark loss were favored by management. Beech (Fagus sylvatica) carried many more microhabitats than silver fir (Abies alba), especially cavities, dendrothelms and bark losses. Fir very scarcely formed dendrothelms. Secondary tree species played an important role by providing cracks and bark losses. The proportion of microhabitat-bearing trees increased dramatically above circumference thresholds of 225 cm for beech and 215 cm for fir. Firs with a circumference of less than 135 cm did not carry microhabitats. In order to conserve microhabitat-providing trees and to increase the volume of dead wood in managed stands, we recommend conserving trees that finish their natural cycle over 10–20% of the surface area.     

Effect of intervention and type of forest management on quality and quantity characteristics of dead wood in managed and reserve forests: A case study

We compared the dead wood (DW) conditions of Cheshmeh-sar forest and Sardab forest with different management history,including reserve forest and harvested forest. The First forest took 100% inventory from all the available DW. Also dead trees were compared interms of species, shape, location and quality of fracture in both forests.Volumes of dead wood in Cheshmeh-sar and Sardab forests were 207.47and 142.74 m3, respectively. Due to this significant difference, impact onthe management level was determined. In Cheshmeh-sar forest, 42% ofdead trees were standing and 58% were fallen type while in Sardab forest 38.6% were standing and 61.4% fallen. But the difference was not statistically significant between them (p = 0.0587). In terms of quality, dead trees of hard, soft and hollow had the highest frequency, respectively.However, 71.5% of DW was seen as hard dead in Cheshmeh-sar forestwhile hard dead trees in Sardab forests were 54.2%. Soft quality degree ofdead trees which formed in Cheshmeh-sar and Sardab forest were calculated as 26.6% and 43.4% respectively. Also 30% of the dead trees of Sardab forest were eradicated while in Cheshmeh-sar this amount was reduced to 12%. Due to this significant difference ((Р=0/018), it is concluded that the type of management and human interference are affecting the quality of dead trees and makes us to think the human interferences could effect on the ecosystem of touched forests.     

Modeling the effects of climate change and management on the dead wood dynamics in boreal forest plantations

The present research examines the joint effects of climate change and management on the dead wood dynamics of the main tree species of the Finnish boreal forests via a forest ecosystem simulator. Tree processes are analyzed in stands subject to multiple biotic and abiotic environmental factors. A special focus is on the implications for biodiversity conservation thereof. Our results predict that in boreal forests, climate change will speed up tree growth and accumulation ending up in a higher stock of dead wood available as habitat for forest-dwelling species, but the accumulation processes will be much smaller in the working landscape than in set-asides. Increased decomposition rates driven by climate change for silver birch and Norway spruce will likely reduce the time the dead wood stock is available for dead wood-associated species. While for silver birch, the decomposition rate will be further increased in set-aside in relation to stands under ordinary management, for Norway spruce, set-asides can counterbalance the enhanced decomposition rate due to climate change thereby permitting a longer persistence of different decay stages of dead wood.     

Carbon dynamics of North American boreal forest after stand replacing wildfire and clearcut logging

Boreal forest carbon (C) storage and sequestration is a critical element for global C management and is largely disturbance driven. The disturbance regime can be natural or anthropogenic with varying intensity and frequency that differ temporally and spatially the boreal forest. The objective of this review was to synthesize the literature on C dynamics of North American boreal forests after most common disturbances, stand replacing wildfire and clearcut logging. Forest ecosystem C is stored in four major pools: live biomass, dead biomass, organic soil horizons, and mineral soil. Carbon cycling among these pools is inter-related and largely determined by disturbance type and time since disturbance. Following a stand replacing disturbance, (1) live biomass increases rapidly leading to the maximal biomass stage, then stabilizes or slightly declines at old-growth or gap dynamics stage at which late-successional tree species dominate the stand; (2) dead woody material carbon generally follows a U-shaped pattern during succession; (3) forest floor carbon increases throughout stand development; and (4) mineral soil carbon appears to be more or less stable throughout stand development. Wildfire and harvesting differ in many ways, fire being more of a chemical and harvesting a mechanical disturbance. Fire consumes forest floor and small live vegetation and foliage, whereas logging removes large stems. Overall, the effects of the two disturbances on C dynamics in boreal forest are poorly understood. There is also a scarcity of literature dealing with C dynamics of plant coarse and fine roots, understory vegetation, small-sized and buried dead material, forest floor, and mineral soil.     

Long-term dynamics of large woody debris in a managed boreal forest stream

Little is known about how past forest management in Sweden influenced the quantity and quality of large woody debris (LWD) in streams. The present study provides information of the long-term dynamics of LWD in a reach of a boreal stream intersecting a managed forest. Dendrochronological methods were used to reconstruct mortality years of the pieces of LWD and the general history of fire and cuttings of the surrounding riparian forest. Today, spruce dominates among the living trees, whereas the LWD is dominated by birch in the forest and by pine in the stream. Fire frequency prior to active fire suppression was similar to values reported from boreal forests. Pine trees were more abundant in the riparian forest before selective logging operations and active fire suppression began in the 1800s. Many of the pieces of LWD found in the stream today died more than 200 years ago and derived from a cohort of pines that generated in the early 1600s. Pine LWD in stream channels is highly resistant to decomposition and can reside for more than 300 years. A substantial amount of the LWD found today in managed forest streams in boreal Sweden most likely derives from the time before extensive human influence and is likely to decrease further in the future. Management of riparian forests to ascertain future supply of long-lived LWD in streams should target to increase the proportion of pine trees.     

Availability of standing trees for large cavity-nesting birds in the eastern boreal forest of Québec,Canada

Large cavity-nesting birds depend on large-diameter trees for suitable nest sites. The increased spatial extent of commercial timber harvesting is modifying forest structure across the land base and may thus compromise the availability of large trees at the landscape scale. In this study, our objectives were to (1) characterize the availability of large living and dead trees in old-growth stands dominated by different tree species and surficial deposits that encompass the range of natural cover types of eastern Québec's boreal forest; (2) analyze the distribution of trees among decay-classes; and (3) compare the availability of large trees in unharvested, remnant, and harvested stands for the entire range of decay-classes. A total of 116 line transects were distributed across unharvested forests, remnant linear forests, and cutblocks in cutover areas. Unharvested forest stands (black spruce [Picea mariana], balsam fir [Abies balsamea]–black spruce, balsam fir–white spruce [Picea glauca] and balsam fir) reflected a gradient of balsam fir dominance. The remnant forests selected were isolated for 5–15 years. Analyses were performed at two diameter cut-off values. Trees with DBH ≥20 cm were considered for availability of total trees whereas trees with DBH ≥30 cm were considered for availability of large trees. Forest stands comprised high proportions of standing dead trees (33% of all stems, 8% were large dead stems). Availability of total and large standing trees increased with the dominance of balsam fir in stands. Forest stands located on thick surficial deposits showed higher densities of large dead trees for every stand type suggesting a higher productivity on those sites. Availability of stems according to decay-classes showed a dome-shaped distribution with higher densities of snags in intermediate decay stages. However, for large stems, black spruce stands showed a significantly lower availability that was consistent across all decay-classes. In linear remnant forests, pure balsam fir stands were absent. Remnant stands thus showed a much lower availability in large trees when compared with unharvested balsam fir stands. Clearcuts had the lowest densities of dead trees across sampled stands. Current even-aged management practices clearly affect availability and recruitment of large trees, therefore forest-dwelling wildlife relying on these structures for breeding is likely to be affected by large-scale harvesting in coniferous boreal forests.     

Spatiotemporal Availability of Dead Wood in Protected Old-growth Forests: A Case Study from Boreal Forests in Eastern Finland

Spatiotemporal patterns of standing and fallen dead trees were examined in two protected Scots pine ( Pinus sylvestris L.)-Norway spruce ( Picea abies (L.) Karst.) forests in eastern Finland (Pahkavaara and Pönttövaara). In Pahkavaara the volume of standing dead trees was 10 m 3 ha -1 and the volume of fallen logs was 66 m 3 ha -1 , and in Pönttövaara the values were 48 m 3 ha -1 and 107 m 3 ha -1 , respectively. The areas differed with respect to the tree species composition, whereas the proportion of different decay stages was similar. Decay stage and dendrochronological analyses revealed the continuity of dead wood formation. The spatial pattern of standing dead trees was usually towards clustered. The volumes of fallen logs showed a spatial autocorrelation up to distances of 10-20 m. The results also suggest that the current amount and diversity of dead wood is rather high, but that forest succession is likely to lead to a less diverse state in the future.     

Changes in quantitative patterns of dead wood in maritime pine plantations over time

Although the volume of dead wood is commonly acknowledged as an indicator of biodiversity in sustainable forest management schemes, only few data are available for plantation forests. To evaluate the volume and qualify the diversity of woody debris along a chronosequence of maritime pine plantations, we sampled downed woody debris, snags and stumps in 143 stands of different ages. To test the hypothesis that the pattern of dead wood accumulation mainly results from silvicultural operations, we developed a predictive model. It combined an empirical growth model evaluating the amount of dead wood produced by successive thinnings with a decay function that estimated the loss of dead wood with time.     

Local spatial pattern in the occurrence of two congeneric wood-decaying fungi in an old-growth boreal forest

Abstract

This paper characterizes spatial patterns in the occurrence of two congeneric wood-decaying fungi (Aphyllophorales: Polyporaceae) in an old-growth boreal forest in eastern Finland. The spatial patterns are used to evaluate indirectly the short-distance dispersal ability of the species. Fomitopsis rosea is a specialist on Picea abies occurring mainly in forests with large amounts of dead wood, whereas Fomitopsis pinicola is a substrate generalist and also abundant in managed forests. Within a forest reserve, a 25 ha study area was divided into 25 m×25 m grid (n=400), and all dead trees and fruiting bodies of the two polypore species were recorded. Spatial patterns were analysed with Spatial Analysis by Distance IndicEs methodology (SADIE). Downed spruce logs were highly aggregated within the study area. After this distribution was accounted for, the spatial pattern of F. pinicola and F. rosea on logs was random. The lack of spatial aggregation suggests that within old-growth forest stands dispersal of the two fungal species is not a limiting factor for their occurrence.