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.     

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.     

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.     

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.     

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.     

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.     

Carbon and nitrogen dynamics along the log bark decomposition continuum in a mesic old-growth boreal forest

Narrowing the uncertainties in carbon (C) and nitrogen (N) dynamics during decomposition of coarse woody debris (CWD) can significantly improve our understanding of forest ecosystem functioning. We examined C, N and pH dynamics in the least studied CWD component—tree bark in a 66-year-long decomposition chronosequence. The relative C concentration decreased by ca. 32% in pine bark, increased by ca. 18% in birch bark and remained stable in spruce and aspen bark. Nitrogen increased in bark of all tree species. In conifer bark, it increased along with epixylic succession. Over 45 years, the relative C/N ratio in bark decreased by 63 and 45% for coniferous and deciduous species, respectively. Bark pH did not change. Due to bark fragmentation, the total C and N amounts in bark of individual logs of aspen, birch, pine and spruce decreased at average rates of 0.03, 0.02, 0.26 and 0.05 year^?1, and 0.02, 0.02, 0.03 and 0.03 year^?1, respectively. At the forest stand level, the total amounts of C and N in log bark were 853 and 21 kg ha^?1 or 11.2 and 45.5% of the C and N amounts stored in downed logs and ca. 2.3–3.8 and 2.2–2.4%, respectively, of total C and N amounts stored in forest litter. In boreal forests, decomposing log bark may act as a long-term source of N for wood-inhabiting communities.     

The effects of burning and dead-wood creation on the diversity of pioneer wood-inhabiting fungi in managed boreal spruce forests

The loss of natural forest habitats due to forestry is the main reason for the decline in boreal forest biodiversity of the Nordic countries of Europe. Ecological rehabilitation may provide means to recover and sustain biodiversity. We analyzed the effects of controlled burning and dead-wood creation (DWC) on the diversity of pioneer wood-inhabiting fungi in managed Norway spruce (Picea abies) forests in southern Finland. Altogether 18 stands were first subjected to a partial cutting with ordinary logging residues in form of cut stumps and treetops left on site. The subsequent rehabilitation treatments consisted of a controlled burning applied in half of the stands and three levels of dead-wood creation (5, 30 and 60 m³ ha⁻¹). The DWC involved creation of logs; felling of whole trees to mimic downed logs formed by natural disturbance processes. Each treatment was replicated three times. Inside each stand, substrates were sampled in two different biotopes; one on mineral soil and one on mineral soil with a thin peat layer. We surveyed the fungal flora on the logs (n = 364) and the ordinary residue stumps (n = 1767) and tops (n = 845) five years after the treatments.When comparing different stands, controlled burning had a significant effect on species composition; certain species were significantly more frequent on substrates in burned stands than in unburned stands, indicating that these species were favored by controlled burning. By contrast, we found no significant effects of DWC levels or biotope on species composition or richness. When comparing different substrates, 99% of the logs hosted at least one species and the occurrence probability of certain species was significantly higher on logs than on ordinary residue stumps and tops. Yet, volume-based rarefaction analyses showed that residues were more species dense than the logs, indicating that ordinary logging residues constitute important resources for many pioneer species.We conclude that controlled burning combined with DWC have strong effects on biodiversity; it modifies the composition of the pioneer wood-inhabiting fungal species found in managed forests and may thereby also influence the further succession and diversity of the secondary fungal flora.     

Stocks of coarse woody debris in old-growth lucidophyllous forests in southwestern Japan

This study quantified the mass and inputs of coarse woody debris (CWD) in two old-growth lucidophyllous forests in southwestern Japan: in a steep slope area at Aya and in a flattish bottomland at Okuchi. CWD mass averaged 36.85 Mg ha⁻¹ with eightfold variations at Aya, and 20.77 Mg ha⁻¹ with more than 40-fold variations at Okuchi. CWD inputs estimated from long-term data on tree mortality averaged 36.76 Mg ha⁻¹ over 16 years at Aya and 44.11 Mg ha⁻¹ over 11 years at Okuchi. In both plots, fallen logs were the major form of CWD mass: 74.4% at Aya and 60.2% at Okuchi. About 19% of CWD was snapped and 7% was uprooted at Aya, and about 34% was snapped and 5.4% was uprooted at Okuchi. The CWD mass differed markedly with topographic conditions in both plots, increasing from valleys up to ridges at Aya and from forest down to a stream at Okuchi. Canopy gaps enhanced CWD mass and inputs in both plots: CWD input under gaps was two to three times that beneath closed canopy. These results imply that typhoons would increase CWD mass and inputs on upper slopes on account of the high aboveground biomass stocks and existence of large-diameter trees.     

Abundance and attributes of wildlife trees and coarse woody debris at three silvicultural systems study areas in the Interior Cedar-Hemlock Zone,British Columbia

Unmanaged cedar (Thuja plicata)-hemlock (Tsuga heterophylla) forests of the northern Interior Wetbelt of British Columbia support standing and dead trees with a variety of structural features that provide habitat for wildlife. We describe the pre-harvest abundance and characteristics of wildlife trees (standing trees with special characteristics that provide habitat for wildlife) and coarse woody debris (CWD) at three silvicultural systems trials in cedar-dominated stands, and the short-term effects of forest harvesting on the abundance and attributes of CWD. The treatments were clearcut, group retention (70% volume removal), group selection (30% volume removal), and unlogged control. We measured standing trees in 75 0.125-ha plots and CWD along 225 24-m transects, using a functional classification system to characterize habitat attributes of trees and logs. CWD assessments were repeated on the same transects after the harvest. The relationship between tree size and occurrence of habitat features was strong for both standing trees and logs. Each of the four major tree species in the study area was associated with specific habitat features that occurred more often in that species than in any other. Large concealed spaces at the bases of trees, providing den sites and escape cover, were associated with hybrid white spruce (Picea engelmannii × glauca). We suggest that these trees had originated on nurse logs that subsequently rotted away; if that supposition is correct, there may be shortages of these structures in future stands that originate from plantations. Forest harvesting had little effect on the volume of CWD, but did affect the decay class distribution, reduce the proportion of pieces having structural habitat attributes, and reduce piece lengths; these effects were generally proportional to the level of harvest removal. Partial-cut silvicultural systems have the potential to mitigate anticipated deficits in large wildlife trees and logs in managed stands, if components of the stand are managed on longer rotations than those planned for timber production alone.     

Diversity of polyporous fungi (Polyporaceae) in northern boreal forests: effects of forest site type and logging intensity

The effects of forest site type and logging intensity on polyporous fungi were studied in subxeric, mesic and herb-rich forests and spruce mires in northern Finland. The species richness of polypores did not follow the fertility gradient of the site types, but was connected with the amount and diversity of coarse woody debris (CWD). The total number of species, and the numbers of indicator and threatened species were equal in subxeric pine forests and in more fertile spruce-dominated stands. The species composition of pine-dominated forests differed conspicuously from that of spruce-dominated site types. The total number of species was not affected by logging intensity, but no virgin forest species or threatened species were found on the sites where the number of cut stumps exceeded 150 stumps ha^?1. Increasing logging intensity decreased the number of polypore observations, indicating reduced substrate availability. The results stress the importance of protecting not only fertile spruce-dominated stands, but also poorer, pine-dominated forests, and sites with high and diverse CWD content.     

Dead wood in clearcuts of semi-natural forests in Estonia: site-type variation, degradation, and the influences of tree retention and slash harvest

Semi-natural forests, which naturally regenerate after timber harvesting, provide distinct opportunities for dead wood (DW) management for biodiversity. We described DW pool and sources of its variation during the first decade after final felling in Estonia, hemiboreal Europe. Depending on forest type, the mean post-harvest volumes of above-ground DW ranged from 70 to 119 m³ ha^?1. Final felling generally did not reduce downed coarse woody debris (CWD) because many sawn logs were left on-site, and soil scarification was rarely used. However, subsequent decay of downed CWD appears to be accelerated due to the increased ground contact of logs, so that even the relatively small inputs from live retention trees observed (5 m³ ha^?1 per decade) can be ecologically significant. While final felling greatly reduced snag abundance, the mortality of retained live trees generally balanced their later losses. The volumes of downed fine woody debris in conventional cutover sites were roughly double that of pre-harvest forests. Slash harvest caused an approximately twofold reduction in downed DW and resulted in CWD volumes that were below mature-forest levels. The results indicate that the habitat quality of cutovers critically depends both on the retention and on the post-harvest management of biological legacies. In Estonia, the necessary improvements include more careful retention of snags in final felling, selecting larger retention trees, focusing slash harvest on the fine debris of common tree species, and providing snags of late-successional tree species.     

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.     

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.     

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.     

Dead wood volume to dead wood carbon: the issue of conversion factors

Requirements for emission reporting under the Kyoto protocol demand an estimate of the dead wood carbon pool in forests. The volume of dead wood consists of coarse woody debris, smaller woody debris and dead roots. The measurement of dead wood volume was included in the most recent National Forest Inventory in Switzerland. To convert dead wood volume into carbon two conversion factors are required: (a) carbon (C) concentration and (b) wood density. So far internationally accepted default values for C concentration (50%) and for wood density (density of alive trees) were used as default values to estimate dead wood carbon, since local measurements were lacking. However, in a field study at 34 sites in Switzerland, the C concentration and density of CWD from Picea abies and Fagus sylvatica of four decay classes were measured recently. The results showed that C concentration in CWD differed significantly between species but did not change due to decay class. The density of CWD decreased significantly with an increase in decay class and it also differed between species. The decrease in CWD density was more pronounced for F. sylvatica than for P. abies. We assessed correlations between climate attributes and CWD density using regression analysis. The modeled densities and measured C concentrations were then expanded with the help of CWD volume data from the NFI3. Spruce CWD and thus spruce CWD carbon is much more abundant in Swiss forests than beech CWD carbon. The majority of spruce CWD is located in the Alps and Pre-Alps. The CWD volume from P. abies was 10 times higher than that from F. sylvatica. Thus, changes in conversion factors for P. abies CWD affected the overall estimate of dead wood carbon in Swiss forests much more than changes in conversion factors for F. sylvatica CWD. Current improvements in CWD conversion factors decreased the estimated amount of spruce CWD carbon by 23.1% and that of beech by 47.6%. The estimated amount of CWD carbon in Swiss forests is decreased by 31%. Since improved estimation methods are currently not applied to smaller woody debris and dead root material, the estimated amount of dead wood carbon is only reduced by 15%. Improving conversion factors for all dead wood fractions would presumably decrease the amount of dead wood carbon by additional 16%.     

Spatial patterns of litter-dwelling taxa in relation to the amounts of coarse woody debris in European temperate deciduous forests

We studied the leaf litter-dwelling fauna of managed deciduous forests and primeval reference sites in Western and Central Europe and addressed the questions if the higher overall species richness close to downed coarse woody debris (CWD) is related to intra-specific or inter-specific aggregation, if the aggregation pattern changes with the amount of CWD on the forest floor, and how much CWD is needed for different taxa. The analysis is based on shelled Gastropoda, Diplopoda/Isopoda, Chilopoda and Coleoptera. Among-sample heterogeneity was lower close to CWD than distant from CWD. This was most pronounced in Diplopoda/Isopoda and Gastropoda. Diplopoda/Isopoda are comparatively mobile and assemblages were already quite homogenous close to CWD at levels above 5 m³ downed deadwood ha⁻¹. Gastropoda have a low mobility, and more than 20 m³ downed deadwood ha⁻¹ is needed for assemblage homogeneity. We further focused on the Gastropoda as sensitive indicators. Enhanced densities and species richness close to CWD were not a simple function of leaf litter weight, thus effects of densities on heterogeneity are not solely driven by leaf litter accumulation close to CWD. In contrast to euryecious litter-dwellers such as the Punctidae, stenecious slow active dispersers such as the Clausiliidae clearly require more than 20 m³ CWD ha⁻¹ for an even distribution. Specialists depending on CWD even seem to have gone extinct in some managed forests. For conserving the litter-dwelling fauna, we propose a target of at least 20 m³ downed CWD ha⁻¹ in already managed forests and rigorous restrictions for deadwood removal from still (almost) pristine systems.     

Patterns of mass,carbon and nitrogen in coarse woody debris in five natural forests in southern China

? Context

Coarse woody debris (CWD, ≥10 cm in diameter) is an important structural and functional component of forests. There are few studies that have estimated the mass and carbon (C) and nitrogen (N) stocks of CWD in subtropical forests. Evergreen broad-leaved forests are distributed widely in subtropical zones in China.

? Aims

This study aimed to evaluate the pools of mass, C and N in CWD in five natural forests of Altingia gracilipes Hemsl., Tsoongiodendron odorum Chun, Castanopsis carlesii (Hemsl.) Hayata, Cinnamomum chekiangense Nakai and Castanopsis fabri Hance in southern China.

? Methods

The mass of CWD was determined using the fixed-area plot method. All types of CWD (logs, snags, stumps and large branches) within the plot were measured. The species, length, diameter and decay class of each piece of CWD were recorded. The C and N pools of CWD were calculated by multiplying the concentrations of C and N by the estimated mass in each forest and decay category.

? Results

Total mass of CWD varied from 16.75 Mg ha^?1 in the C. fabri forest to 40.60 Mg ha^?1 in the A. gracilipes forest; of this CWD, the log contribution ranged from 54.75 to 94.86 %. The largest CWD (≥60 cm diameter) was found only in the A. gracilipes forest. CWD in the 40–60 cm size class represented above 65 % of total mass, while most of CWD accumulations in the C. carlesii, C. chekiangense and C. fabri forests were composed of pieces with diameter less than 40 cm. The A. gracilipes, T. odorum, C. carlesii and C. chekiangense forests contained the full decay classes (from 1 to 5 classes) of CWD. In the C. fabri forest, the CWD in decay classes 2–3 accounted for about 90 % of the total CWD mass. Increasing N concentrations and decreasing densities, C concentrations, and C:N ratios were found with stage of decay. Linear regression showed a strong correlation between the density and C:N ratio (R ²?=?0.821). CWD C-stock ranged from 7.62 to 17.74 Mg ha^?1, while the N stock varied from 85.05 to 204.49 kg ha^?1. The highest overall pools of C and N in CWD were noted in the A. gracilipes forest.

? Conclusion

Differences among five forests can be attributed mainly to characteristics of the tree species. It is very important to preserve the current natural evergreen broad-leaved forest and maintain the structural and functional integrity of CWD.     

缙云山常绿阔叶林粗木质残体储量及特征

【目的】实地调查缙云山常绿阔叶林粗木质残体(CWD)的储量和特征,分析其影响因素,为深入认识常绿阔叶林生态系统中与粗木质残体相关的物质循环等关键生态过程提供理论依据,并为全球碳汇及相关学科的研究提供基础数据。【方法】以缙云山常绿阔叶林内已建立的2块0.5 hm^2样地为对象,调查并分析样地CWD的储量、类型组成、分解等级、径级分布格局以及地形对CWD储量的影响。【结果】缙云山常绿阔叶林CWD储量为38.42 t·hm^-2,其中倒木、枯立木、大枯枝和木桩储量分别为27.70,4.91,2.91和2.90 t·hm^-2,倒木是CWD的主要组成部分; CWD的树种组成与群落优势树种的组成相似,栲占CWD总量的88.44%;从径级分布来看,直径>35 cm的倒木和枯立木是CWD的主体; CWD主要处于分解中后期,中级分解和高级分解CWD的比例分别为63.20%和23.01%;地形对CWD的分布具有显著影响,坡面和山脊CWD储量高于沟谷(P<0.01)。【结论】缙云山常绿阔叶林CWD储量较大,处于该类森林系统的前列; CWD主要以处于分解中后期的优势种大径级倒木和枯立木为主,并主要分布在坡面和沟谷。在森林管理中应降低对CWD的人为干扰和去除,维持森林中枯立木、倒木、大枯枝和树桩等组分的自然状态。     

Stocks and dynamics of soil organic carbon and coarse woody debris in three managed and unmanaged temperate forests

The effect of forest conservation on the organic carbon (C) stock of temperate forest soils is hardly investigated. Coarse woody debris (CWD) represents an important C reservoir in unmanaged forests and potential source of C input to soils. Here, we compared aboveground CWD and soil C stocks at the stand level of three unmanaged and three adjacent managed forests in different geological and climatic regions of Bavaria, Germany. CWD accumulated over 40–100 years and yielded C stocks of 11 Mg C ha^?1 in the unmanaged spruce forest and 23 and 30 Mg C ha^?1 in the two unmanaged beech–oak forests. C stocks of the organic layer were smaller in the beech–oak forests (8 and 19 Mg C ha^?1) and greater in the spruce forest (36 Mg C ha^?1) than the C stock of CWD. Elevated aboveground CWD stocks did not coincide with greater C stocks in the organic layers and the mineral soils of the unmanaged forests. However, radiocarbon signatures of the O _e and O _a horizons differed among unmanaged and managed beech–oak forests. We attributed these differences to partly faster turnover of organic C, stimulated by greater CWD input in the unmanaged forest. Alternatively, the slower turnover of organic C in the managed forests resulted from lower litter quality following thinning or different tree species composition. Radiocarbon signatures of water-extractable dissolved organic carbon (DOC) from the top mineral soils point to CWD as potent DOC source. Our results suggest that 40–100 years of forest protection is too short to generate significant changes in C stocks and radiocarbon signatures of forest soils at the stand level.