Concept and Classification of Coarse Woody Debris in Forest Ecosystems

Coarse woody debris (CWD) is generally considered as dead woody materials in various stages of decomposition,including sound and rotting logs,snags,and large branches.CWD is an important functional and structural component of forested ecosystems and plays an important role in nutrient cycling,long-term carbon storage,tree regeneration,and maintenance of heterogeneous environmental and biological diversity.However,the definition and classification of CWD have been the subject of a long debate in forest ecology.CWD has not been precisely defined.Recently,with the rapid development of landscape ecology in CWD,the USDA Forest Service and the Long Term Ecological Research (LTER)have provided a standardized definition and classification for CWD,which makes data comparison in landscape scale possible.Important characteristics of their definition include:(1) a minimum diameter (or an equivalent crosssection) of CWD≥10 cm at the widest point (the woody debris with a diameter from 1 to 10 cm should be defined as fine woody debris,and the rest is litterfall);and (2) sound and rotting logs,snags,stumps,and large branches (located above the soil),and coarse root debris (larger than 1 cm in diameter).This classification has greatly facilitated CWD studies.Therefore,it has been widely applied in some countries (particularly in North America).However,this classification has long been a source of confusion for forest ecologists in China.Furthermore,different definitions and criteria are still adopted in individual studies,which makes the interpretation and generalization of their work difficult.This article reviewed recent progress in classifying CWD,with an emphasis on introducing the classification system of the USDA Forest Service and the LTER.It is expected that this review will help facilitate the development of standardized definition and classification suitable to forest ecosystems in China.     

Concept and Classification of Coarse Woody Debris in Forest Ecosystems

Coarse woody debris (CWD) is generally considered as dead woody materials in various stages of decomposition, including sound and rotting logs, snags, and large branches. CWD is an important functional and structural component of forested ecosystems and plays an important role in nutrient cycling, long-term carbon storage, tree regeneration, and maintenance of heterogeneous environmental and biological diversity. However, the definition and classification of CWD have been the subject of a long debate in forest ecology. CWD has not been precisely defined. Recently, with the rapid development of landscape ecology in CWD, the USDA Forest Service and the Long Term Ecological Research (LTER) have provided a standardized definition and classification for CWD, which makes data comparison in landscape scale possible. Important characteristics of their definition include: (1) a minimum diameter (or an equivalent cross-section) of CWD ≥10 cm at the widest point (the woody debris with a diameter from 1 to 10 cm should be defined as fine woody debris, and the rest is litterfall); and (2) sound and rotting logs, snags, stumps, and large branches (located above the soil), and coarse root debris (larger than 1 cm in diameter). This classification has greatly facilitated CWD studies. Therefore, it has been widely applied in some countries (particularly in North America). However, this classification has long been a source of confusion for forest ecologists in China. Furthermore, different definitions and criteria are still adopted in individual studies, which makes the interpretation and generalization of their work difficult. This article reviewed recent progress in classifying CWD, with an emphasis on introducing the classification system of the USDA Forest Service and the LTER. It is expected that this review will help facilitate the development of standardized definition and classification suitable to forest ecosystems in China. Translated from Acta Ecologica Sinica, 2005, 25(1) (in Chinese)     

Coarse Woody Debris Quantity and Distribution in Central European Streams

Summarized here are ten investigations concerning the volume of coarse woody debris (CWD) in Central European streams. Altogether, 69 stream sections were examined ranging from Northern German lowland streams to brooks in alpine regions. Most of the study streams are according to Central European standards quasi‐natural and are bordered by deciduous forest. The geometric mean of CWD volume related to stream length is 1.44 m³ /100 meter reach. Related to stream bottom area, the geometric mean of CWD volume is 0.202 m³ /100 m² . The mean number of logs (≥10 cm diameter) is 12.5 logs/100 meter reach, and 3.01/100 m² bottom area (geometric means). Regarding only quasi‐natural stream sections (riparian forest currently unmanaged and no removal of CWD for at least 10 years), the geometric mean of CWD standing stock is 0.45 m³ /100 m² for lowland streams, 0.38 m³/100 m² for streams in lower mountainous areas and 0.02 m³ /100 m² for alpine floodplains. From the distribution of size classes and comparison with other studies it is likely, that the current CWD standing stock is considerably less than the potential amount of CWD. For centuries all of the streams have been influenced by man. Historic alterations of the stream, its floodplain and the riparian vegetation may still affect CWD supply and standing stock. We conclude that virtually all streams in Central Europe are highly altered with respect to the amount of CWD, and that the importance of CWD is under‐represented in recent assessment principles for streams in Germany.     

鼎湖山锥栗粗木质残体的分解和元素动态

选取锥栗(Castanopsischinensis)粗木质残体(coarsewoodydebris,CWD)的3个径级(径级1-3分别为5-10cm,10-20cm和20-30cm),并且将每个径级的锥栗粗木质残体分为Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅴ5个分解阶段。通过测定锥栗粗木质残体3个径级的Ⅰ-Ⅲ阶段的7种化学元素(C、N、P、K、Ca、Na、Mg)的浓度和CWD密度变化来研究其分解过程中的元素动态、分解速率及其分解过程中基质质量的变化。到分解阶段Ⅲ时,3个径级的重量与原来相比分别损失了36%、48%和43%。元素N、P、Mg、Ca、Na的浓度升高。元素N的累积可能和锥栗粗木质残体中寄生着固氮细菌和真菌有关。元素P、Mg、Ca、Na的浓度升高则可能是由于这4个元素因淋溶流失的速度小于锥栗粗木质残体质量的损失速度,造成元素积聚,元素C、K的浓度降低。N/P比值是较好的分解指示指标。虽然存在元素的净释放,但是由于C和其它主要元素的释放速率较慢,因而锥栗粗木质残体是森林中重要的C库和长期的元素来源。     

Effects of Forest Type and Disturbance on Diversity of Coarse Woody Debris in Boreal Forest

Coarse woody debris (CWD) volume and diversity are vital attributes of forest ecosystems. However, despite their importance, their long-term dynamics associated with fire- or logging-origin and overstory type have not been examined in boreal forest. We hypothesize that (1) CWD compositional diversity increases with stand development whereas CWD volume follows a U-shaped pattern. Furthermore, we attempted to test if (2) CWD volume and compositional diversity converge for postlogged and postfire stands through stand development, and (3) mixedwoods have more CWD volume and greater compositional diversity than conifer or broadleaf overstory types. We sampled 72 stands ranging in age from 7 to 201 years in fire-origin stands and 7–31 years in managed stands with conifer, mixedwood, and broadleaf overstory types in central boreal Canada. For fire-origin stands, snag volume was 100–260 m³/ha in 7-year-old stands, 5–20 m³/ha in 25-year-old stands, and 25–60 m³/ha in older stands; downed woody debris (DWD) volume decreased from 7 to 72–90 year-old stands, increased in 124- to 139-year-old stands, then either decreased or increased in 201-year-old stands depending on overstory type. CWD diversity increased from 25 to 124–139 year-old and plateaued, but in 7-year-old stands, CWD diversity was as high as that in the 124 and up year-old age classes. Logging resulted in a smaller amount and lower size variability of CWD in 7-year-old stands, with a larger portion being fast-decomposing Betula papyrifera. Most CWD characteristics had not converged by approximately 30 years since disturbance between the two stand origins. More diverse CWD occurred in mixedwoods, but conifer stands contained the greatest CWD volume except in 7 year-old postfire stands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. B. W. Brassard collected and analyzed data and wrote the paper. H. Y. H. Chen conceived and designed the study, analyzed data, and critiqued earlier drafts of the paper.     

Preliminary Studies on the Litterfall and Coarse Woody Debris in Mid-Mountain Humid Evergreen Broad-Leaved Forest in Ailao Mountains

The composition and amount of the litterfall and coarse woody debris (CWD), their time and space distribution and the element return were studied in mid-mountain humid ev-ergreen broad-leaved forest in Ailao Mountain in Yunnan, China. The average litterfall (leaves, wood (< 2.5 cm diameter), flower, fruit and mixed matter) was (6.77± 1.43) t/(hm2 · a) while the large wood litterfall () 2.5 cm diameter) was (0.45±0.18) t/(hm2 · a). There were two marked peaks of annual litterfalh the main one from April to May and the second one from October to November. The standing crop of CWD in the forest was 98.46 t/hm2, in which logs made up to 85.8% of the total. As the CWD was not well-distributed in the forest, it increased the diversities of structure and site in the forest ecosystem. The total storage of C, N, P, K, Ca, Mg, Mn, A1 and Fe in the CWD was 48699 t/hm2, but return of elements to the forest ground in the litterfall only made up 7.77% of the nutrient storage of the CWD.     

Coarse Woody Debris following Fire and Logging in Wyoming Lodgepole Pine Forests

The accumulation and decomposition of coarse woody debris (CWD) are processes that affect habitat, soil structure and organic matter inputs, and energy and nutrient flows in forest ecosystems. Natural disturbances such as fires typically produce large quantities of CWD as trees fall and break, whereas human disturbances such as timber harvesting remove much of the CWD. Our objective was to compare the amount of CWD removed and left behind after clear-cutting to the amount consumed and left behind after natural fires in Rocky Mountain lodgepole pine. The masses of fallen logs, dead-standing trees, stumps, and root crowns more than 7.5 cm in diameter were estimated in clear-cut and intact lodgepole pine forests in Wyoming and compared to estimates made in burned and unburned stands in Yellowstone National Park (YNP), where no timber harvesting has occurred. Estimates of downed CWD consumed or converted to charcoal during an intense crown fire were also made in YNP. No significant differences in biomass of downed CWD more than 7.5 cm in diameter were detected between burned stands and those following a single clear-cut. However, the total mass of downed CWD plus the mass of snags that will become CWD was nearly twice as high in burned stands than in clear-cuts. In YNP, approximately 8% of the downed CWD was consumed by fire and an additional 8% was converted to charcoal, for an estimated loss of about 16%. In contrast, approximately four times more wood (70%) was removed by clear-cutting. Considering all CWD more than 7.5 cm in diameter that was either still present in the stand or removed by harvesting, slash treatment, or burning, clear-cut stands lost an average of 80 Mg ha⁻¹ whereas stands that burned gained an average of 95 Mg ha⁻¹. Some CWD remains as slash and stumps left behind after harvesting, but stands subjected to repeated harvesting will have forest floor and surface soil characteristics that are beyond the historic range of variability of naturally developing stands. Received 16 November 1999; Accepted 31 May 2000.     

Current Loads of Coarse Woody Debris on Southeastern Australian Floodplains: Evaluation of Change and Implications for Restoration

We evaluated the status of coarse woody debris (CWD, fallen wood) on floodplains of the southern Murray‐Darling basin of southeastern Australia. The floodplains are dominated floristically by the river red gum Eucalyptus camaldulensis. Aerial survey techniques were used to estimate the amounts of woody debris within 200 m of the channels along 2,442 km of 11 rivers of the system, including the Murray and Darling Rivers and the Darling Anabranch. Aerially based indices were converted into wood volumes by using ground‐truthing at a selection of sites; there was a strong correlation between index values and measured wood volume densities. For thickly forested sites such as Barmah, Gunbower Island, and the Ovens floodplains, the aerial method was not useful, so ground measurements at randomly positioned sites within the forests were used. Volumes were translated into mass by using conversion factors drawn from the literature. We estimated that total tonnage on approximately 221,000 ha of floodplain forests was 4.175 ± 0.579 × 10⁶ tonne. In the larger forested blocks (>7,000 ha), mean wood densities ranged between approximately 12 tonne/ha on the lower Goulburn up to approximately 24 tonne/ha at Barmah State Forest. The area‐weighted mean for the entire area was approximately 19 tonne/ha. A main purpose of the research was to place these figures into an historical perspective to evaluate implications for restoration. A thorough search of historical documentation revealed that there are no extant data upon which to estimate pre‐European settlement levels. We used information from an apparently undisturbed “unmanaged” site in the Millewa forests of southern New South Wales as a basis. Wood density there corresponded to a mean figure of 125 tonne/ha wood‐mass density. By using this figure we estimate that CWD levels on the southern Murray‐Darling basin may be of the order of 15% of pre‐European settlement levels. Full restoration of the 221,000 ha surveyed would require 23.5 ± 0.579 × 10⁶ tonne, which is equivalent to about 600,000 mature (1 m diameter at breast height) river red gum trees or the amount of timber derived from clear felling about 115,000 ha of river red gum forest at current stocking levels. We discuss the implications of this massive deficit and possible short‐ and long‐term solutions.     

Carbon distribution of a well- and poorly-drained black spruce fire chronosequence

The objective of this study was to quantify carbon (C) distribution for boreal black spruce (Picea mariana (Mill.) BSP) stands comprising a fire chronosequence in northern Manitoba, Canada. The experimental design included seven well‐drained (dry) and seven poorly‐drained (wet) stands that burned between 1998 and 1850. Vegetation C pools (above‐ground + below‐ground) steadily increased from 1.3 to 83.3 t C ha^?1 for the dry chronosequence, and from 0.6 to 37.4 t C ha^?1 for the wet chronosequence. The detritus C pools (woody debris + forest floor) varied from 10.3 to 96.0 t C ha^?1 and from 12.6 to 77.4 t C ha^?1 for the dry and wet chronosequence, respectively. Overstorey biomass, mean annual biomass increment (MAI), woody debris mass, and litterfall were significantly greater (α = 0.05) for the dry stands than for the wet stands, but the bryophyte, understorey, and forest floor C pools were significantly less for the dry than for the wet stands. The root mass ratio decreased with stand age until 37 years after fire, was fairly constant thereafter, and was not significantly affected by soil drainage. The C pools of the overstorey and bryophyte tended to increase with stand age. Foliage biomass, litterfall, and MAI (for the dry stands) peaked at 71 years after fire and declined in the oldest stands. The results from this study illustrate that the effects of disturbance and edaphic conditions must be accounted for in boreal forest C inventories and C models. The appropriateness of using chronosequences to examine effects of wildfire on ecosystem C distribution is discussed.     

Dissolved Organic Carbon in Alaskan Boreal Forest: Sources, Chemical Characteristics, and Biodegradability

The fate of terrestrially-derived dissolved organic carbon (DOC) is important to carbon (C) cycling in both terrestrial and aquatic environments, and recent evidence suggests that climate warming is influencing DOC dynamics in northern ecosystems. To understand what determines the fate of terrestrial DOC, it is essential to quantify the chemical nature and potential biodegradability of this DOC. We examined DOC chemical characteristics and biodegradability collected from soil pore waters and dominant vegetation species in four boreal black spruce forest sites in Alaska spanning a range of hydrologic regimes and permafrost extents (Well Drained, Moderately Well Drained, Poorly Drained, and Thermokarst Wetlands). DOC chemistry was characterized using fractionation, UV–Vis absorbance, and fluorescence measurements. Potential biodegradability was assessed by incubating the samples and measuring CO₂ production over 1 month. Soil pore water DOC from all sites was dominated by hydrophobic acids and was highly aromatic, whereas the chemical composition of vegetation leachate DOC varied significantly with species. There was no seasonal variability in soil pore water DOC chemical characteristics or biodegradability; however, DOC collected from the Poorly Drained site was significantly less biodegradable than DOC from the other three sites (6% loss vs. 13–15% loss). The biodegradability of vegetation-derived DOC ranged from 10 to 90% loss, and was strongly correlated with hydrophilic DOC content. Vegetation such as Sphagnum moss and feathermosses yielded DOC that was quickly metabolized and respired. In contrast, the DOC leached from vegetation such as black spruce was moderately recalcitrant. Changes in DOC chemical characteristics that occurred during microbial metabolism of DOC were quantified using fractionation and fluorescence. The chemical characteristics and biodegradability of DOC in soil pore waters were most similar to the moderately recalcitrant vegetation leachates, and to the microbially altered DOC from all vegetation leachates.     

Changing sources of soil respiration with time since fire in a boreal forest

Radiocarbon signatures (Δ¹⁴C) of carbon dioxide (CO₂) provide a measure of the age of C being decomposed by microbes or respired by living plants. Over a 2‐year period, we measured Δ¹⁴C of soil respiration and soil CO₂ in boreal forest sites in Canada, which varied primarily in the amount of time since the last stand‐replacing fire. Comparing bulk respiration Δ¹⁴C with Δ¹⁴C of CO₂ evolved in incubations of heterotrophic (decomposing organic horizons) and autotrophic (root and moss) components allowed us to estimate the relative contributions of O horizon decomposition vs. plant sources. Although soil respiration fluxes did not vary greatly, differences in Δ¹⁴C of respired CO₂ indicated marked variation in respiration sources in space and time. The ¹⁴C signature of respired CO₂ respired from O horizon decomposition depended on the age of C substrates. These varied with time since fire, but consistently had Δ¹⁴C greater (averaging ～120‰) than autotrophic respiration. The Δ¹⁴C of autotrophically respired CO₂ in young stands equaled those expected for recent photosynthetic products (70‰ in 2003, 64‰ in 2004). CO₂ respired by black spruce roots in stands >40 years old had Δ¹⁴C up to 30‰ higher than recent photosynthates, indicating a significant contribution of C stored at least several years in plants. Decomposition of O horizon organic matter made up 20% or less of soil respiration in the younger (<40 years since fire) stands, increasing to ～50% in mature stands. This is a minimum for total heterotrophic contribution, since mineral soil CO₂ had Δ¹⁴C close to or less than those we have assigned to autotrophic respiration. Decomposition of old organic matter in mineral soils clearly contributed to soil respiration in younger stands in 2003, a very dry year, when Δ¹⁴C of soil respiration in younger successional stands dropped below those of the atmospheric CO₂.     

天宝岩自然保护区长苞铁杉混交林粗木质残体蓄水能力研究

选择天宝岩国家级自然保护区内4种不同长苞铁杉混交林(A~D)粗木质残体(CWD)为研究对象,采用自然风干法与室内浸泡法对不同类型(倒木、枯立木、树桩)和不同腐烂程度(Ⅰ~Ⅴ级,腐烂由轻到重)CWD的蓄水能力进行测定分析和对比研究。结果显示:(1)CWD有效拦蓄量在不同长苞铁杉混交林下表现为:长苞铁杉+石栎+马尾松混交林(B,14.21t/hm2)长苞铁杉+甜槠+青冈混交林(C,7.78t/hm2)长苞铁杉+青冈+深山含笑混交林(D,6.99t/hm2)长苞铁杉+猴头杜鹃混交林(A,4.74t/hm2),而在各CWD类型下表现为:倒木(19.579t/hm2)枯立木(8.494t/hm2)树桩(5.661t/hm2)。(2)各长苞铁杉混交林CWD的蓄水量和蓄水速率在蓄水过程中都表现为BDCA,而在脱水过程中都表现为BCAD。(3)不同腐烂等级的CWD按其蓄水量、蓄水速率和脱水速率大小排列皆为:Ⅱ级Ⅵ级Ⅲ级Ⅴ级Ⅰ级。研究表明:CWD蓄水量的大小差异与群落类型、腐烂等级、CWD本身持水性等因素有关,其蓄水和脱水的速率与其持水量、蓄积量以及腐烂等级之间也存在密切的联系;CWD的自然持水率随腐烂等级增加呈增加趋势,而且腐烂等级越高CWD蓄水速率变化也越明显,CWD腐烂等级对其蓄水能力影响较大;倒木具有较好的持水能力,并在蓄积量上远大于树桩和枯立木,对降雨有较强的实际拦截能力。     

Rates of lateral channel migration along the Mala Panew River (southern Poland) based on dating riparian trees and Coarse Woody Debris

Two methods of estimating the lateral migration of a river channel have been proposed. The first method is based on Coarse Woody Debris (CWD) dating. The terraces of the Mala Panew River are mainly covered with plantations of Pinus sylvestris where individual trees grow at equal distances to each other. During times of high discharges trees fall onto the riverbed providing information on the extent of flood plain erosion. The ages of CWD and the surface of the eroded flood plain provide an estimation of the rate of lateral migration. The erosion rates measured at two sites in the Mala Panew River were between 0.24 and 0.36 m/year.Another way of reconstructing the rate of lateral migration is by dating trees growing on different-aged sandy meander bars. These levels are primarily covered with Alnus glutinosa and Alnus incana. The oldest trees growing on each level give information about the minimum age of that level, which allows us to reconstruct the rate of lateral migration. The lateral migration of the channel has also been estimated dating the oldest trees growing on mid-channel islands separated from the lateral banks. The values obtained for 10 sites of the Mala Panew channel oscillate between 0.07 and 1.83 m/year.Tree ring analyses also allow us to determine the impact of individual high discharges on the lateral migration rate of the Mała Panew channel. The lateral migration of the channel was most rapid in the years 1953–57 and 1966–68 as well as during the extraordinary flood in 1997.     

The Imprint of Land-use History: Patterns of Carbon and Nitrogen in Downed Woody Debris at the Harvard Forest

Few data sets have characterized carbon (C) and nitrogen (N) pools in woody debris at sites where other aspects of C and N cycling are studied and histories of land use and disturbance are well documented. We quantified pools of mass, C, and N in fine and coarse woody debris (CWD) in two contrasting stands: a 73-year-old red pine plantation on abandoned agricultural land and a naturally regenerated deciduous forest that has experienced several disturbances in the past 150 years. Masses of downed woody debris amounted to 40.0 Mg ha⁻¹ in the coniferous stand and 26.9 Mg ha⁻¹ in the deciduous forest (20.4 and 13.8 Mg C ha⁻¹, respectively). Concentrations of N were higher and C:N ratios were lower in the deciduous forest compared to the coniferous. Pools of N amounted to 146 kg N ha⁻¹ in the coniferous stand and 155 kg N ha⁻¹ in the deciduous forest; both are larger than previously published pools of N in woody debris of temperate forests. Woody detritus buried in O horizons was minimal in these forests, contrary to previous findings in forests of New England. Differences in the patterns of mass, C, and N in size and decay classes of woody debris were related to stand histories. In the naturally regenerated deciduous forest, detritus was distributed across all size categories, and most CWD mass and N was present in the most advanced decay stages. In the coniferous plantation, nearly all of the CWD mass was present in the smallest size class (less than 25 cm diameter), and a recognizable cohort of decayed stems was evident from the stem-exclusion phase of this even-aged stand. These results indicate that heterogeneities in site histories should be explicitly included when biogeochemical process models are used to scale C and N stocks in woody debris to landscapes and regions. Received 27 April 2001; accepted 4 January 2002.     

The Effect of a Copper Smelter Emissions on the Stock and Decomposition of Coarse Woody Debris in Spruce and Fir Woodlands

We have studied the share of coarse woody debris (CWD) reserves at different decay classes in the spruce and fir woodland within the impact area of aerial pollution from the Middle Ural Copper Smelter (Revda, Sverdlovsk oblast). Control and impact areas slightly differ in total reserves and number of trunks of CWD (sum of standing and fallen dead wood). However, the number of CWD tends to grow in proximity to the plant. The mechanisms involved in CWD-reserve formation differ between impact and control sites. A larger number of relatively thin trunks prevail in CWD reserves of impact sites when compared to the lower number of thick trunks at control sites. The CWD share of 30% in the total number of dead and living trees did not differ across pollution loads. However, the share of CWD reserves in total stock is 1.9 times higher near the plant than at the control site. The share of logs at the initial stages of decomposition (first and second decay classes) is 3.2 times higher in terms of CWD number and 4.2 times higher in terms of CWD reserves than at the control sites. This points to the strong inhibition of CWD decomposition. The pattern of decay classes of all sizes of fallen trees significantly differs in volume across pollution zones.     

Interactive Effects of Fire, Soil Climate, and Moss on CO2 Fluxes in Black Spruce Ecosystems of Interior Alaska

Fire is an important control on the carbon (C) balance of the boreal forest region. Here, we present findings from two complementary studies that examine how fire modifies soil organic matter properties, and how these modifications influence rates of decomposition and C exchange in black spruce (Picea mariana) ecosystems of interior Alaska. First, we used laboratory incubations to explore soil temperature, moisture, and vegetation effects on CO₂ and DOC production rates in burned and unburned soils from three study regions in interior Alaska. Second, at one of the study regions used in the incubation experiments, we conducted intensive field measurements of net ecosystem exchange (NEE) and ecosystem respiration (ER) across an unreplicated factorial design of burning (2 year post-fire versus unburned sites) and drainage class (upland forest versus peatland sites). Our laboratory study showed that burning reduced the sensitivity of decomposition to increased temperature, most likely by inducing moisture or substrate quality limitations on decomposition rates. Burning also reduced the decomposability of Sphagnum-derived organic matter, increased the hydrophobicity of feather moss-derived organic matter, and increased the ratio of dissolved organic carbon (DOC) to total dissolved nitrogen (TDN) in both the upland and peatland sites. At the ecosystem scale, our field measurements indicate that the surface organic soil was generally wetter in burned than in unburned sites, whereas soil temperature was not different between the burned and unburned sites. Analysis of variance results showed that ER varied with soil drainage class but not by burn status, averaging 0.9 ± 0.1 and 1.4 ± 0.1 g C m⁻² d⁻¹ in the upland and peatland sites, respectively. However, a more complex general linear model showed that ER was controlled by an interaction between soil temperature, moisture, and burn status, and in general was less variable over time in the burned than in the unburned sites. Together, findings from these studies across different spatial scales suggest that although fire can create some soil climate conditions more conducive to rapid decomposition, rates of C release from soils may be constrained following fire by changes in moisture and/or substrate quality that impede rates of decomposition. Author contributions: JAO: performed research, analyzed data, contributed new methods, wrote the paper; MRT: designed laboratory study, performed research, analyzed data; JWH: designed field study, performed research; KLM: performed research; LEP: performed research, contributed new method; GS: performed research; JCN: performed research.     

Home Range Size and Micro‐habitat Density Requirements of Egernia napoleonis: Implications for Restored Jarrah Forest of South Western Australia

Compared to natural forests, coarse woody debris (CWD) is typically scarce in restored forests due to the long time it takes to develop naturally. In post‐mining restored forests in the Jarrah forest of south western Australia, CWD is returned at densities of one log pile per hectare. We tested the adequacy of these densities for meeting the micro‐habitat requirements of Napoleon's skink (Egernia napoleonis), a species rarely found within restored sites. Home range size and overlap, and micro‐habitat densities used by skinks, were measured by radio‐tracking 12 individuals in natural, unmined forest. Napoleon's skinks had small home ranges (0.08 ± 0.02 ha), based on 8 individuals with sufficient fixes. All skinks overlapped in home ranges, with average overlaps of 43.5 ± 8.6%. Ten of the 12 skinks shared micro‐habitats and 4 shared them simultaneously, which indicates some social tolerance. This will influence as to how many micro‐habitats are required. Micro‐habitats were used at high densities: logs at 49.2 ± 8.8 ha^?1 and woody debris piles at 12.4 ± 4.8 ha^?1. Based on these densities, it is recommended that CWD is returned to restored forests at densities of 60 ha^?1, which should provide sufficient micro‐habitats for multiple skinks. Due to the infeasibility of returning such CWD densities across large areas of restored forest, CWD could be preferentially returned as patches, large enough for numerous home ranges, adjacent to unmined forest, or as corridors between unmined forest. These recommendations for returning micro‐habitats should be tested for effectiveness in encouraging recolonization of restored forest by Napoleon's skink and other species.     

Demography of Black Bears in Hunted and Unhunted Areas of the Boreal Forest of Ontario

Abstract: We estimated relative density, survival, and reproduction of American black bears (Ursus americanus) from capture-recapture and telemetry data collected from 1989 to 1999 in the unhunted Chapleau Crown Game Preserve (CCGP) and nearby hunted areas in the boreal forest of Ontario, Canada. We tested for combinations of effects of age class, sex, year, years of food shortage, encumbrance status, and residency (on or off the Game Preserve) on vital rates. Results from live captures, remote captures, and bait-station hit rates indicated that density was highest inside CCGP. Total survival of adult females, subadults, and cubs were similar among residents and nonresidents of CCGP, but yearling survival was lower among CCGP residents. Adult females were approximately twice as likely to die and nearly 10 times as likely to be cannibalized (risk ratio [RR] = 9.62, 95% CI = 2.088–44.29) while encumbered with cubs of the year. Nonresidents of CCGP had greater risk of being harvested (RR = 4.00, 95% CI = 1.19–13.46) but similar risk of being cannibalized (RR = 0.875, 95% CI = 0.300–2.55) relative to CCGP residents, suggesting that harvest mortality was additive to other forms of mortality. Residents of CCGP had older ages at primiparity and lower litter-production rates than bears resident in hunted areas. Few litters were produced in years following food shortages, but litter size was unaffected. We recommend that managers allow for additive harvest mortality and reduced survival of bears encumbered with cubs of the year, and we caution that assuming density-compensatory increases in cub production could optimistically bias estimates of population growth.     

Relative roles of termites and saprotrophic microbes as drivers of wood decay: A wood block test

Deadwood in tropical ecosystems represents an important but poorly studied carbon (C) pool. Biologically mediated decay of this pool occurs by both saprotrophic microbes and macro‐invertebrates, such as termites. The activity of these decay agents is influenced by abiotic conditions, especially water availability in tropical systems. While saprotrophic microbial activity is directly controlled by moisture, termites employ various morphological and behavioural modifications that should allow for continued activity in dry conditions. We therefore hypothesized that the relative role of termites would be enhanced in the dry season and a dry compared to a wet site. We deployed a novel wood bait (Pinus radiata) at two sites (rainforest and savanna), with or without access holes cut into termite‐excluding mesh. Mass loss from wood baits was measured after a dry season and after a full dry/wet annual cycle. Mass loss was higher at the rainforest site, demonstrating the overall role of moisture in driving wood decay. Counter to expectations, we found no evidence that the relative role of termites was higher at the dry site, nor during the dry season. However, the prevalence of termites was higher in the savanna compared to the rainforest. While termites clearly impact wood decay, these findings indicate that the relative importance of termites in the fate of deadwood may not reflect their mere presence within and across ecosystems. If moisture availability shifts under climate change, our results suggest similar functional responses between termites and saprotrophic microbes in driving C loss from deadwood.