Decay rates of above- and below-ground coarse woody debris of common tree species in New Zealand’s natural forest

Coarse woody debris decay rates for seven indigenous tree species in the central North Island of New Zealand

Mark O. Kimberley, Peter N. Beets and Thomas S.H. Paul, Scion, Rotorua, New Zealand Background/Question/Methods New Zealand uses a national forest inventory to assess carbon in live and dead pools in its natural forest. Coarse woody debris (CWD) carbon is estimated by multiplying volumes of dead stems and fallen pieces with wood and bark density, a density modifier based on a 5-level decay class scale, and the carbon fraction. Two cycles of the inventory have been completed with both providing CWD carbon estimates of approximately 20 t C ha -1 indicating the CWD pool is not changing over time. Independently, exponential decay constants have been determined for common tree species from measurements of fallen stems. However, predicted annual decay loses from the CWD pool based on these decay constants are substantially lower than inventory estimates of tree mortality. This discrepancy provides evidence that the inventory substantially underestimates the CWD pool. In the current study, decay rates were estimated using an alternative approach based on the proportion of matched pieces of CWD transferring across decay classes between inventory measurements. The objective of the study was to compare inventory estimates of CWD carbon stock changes with stock changes predicted using decay constants and measured mortality, and identify likely causes of discrepancies. Results/Conclusions The half-life or time for CWD to lose 50% dry weight in New Zealand’s natural forest was estimated from the proportion of matched pieces transferring across decay classes between inventory assessments, and the quantity of stocks in each decay class. The volume-weighted average half-life was 25.8 years, almost identical to the estimate of 25.5 years based on decay constants derived from measurements of fallen stems obtained independently of the inventory. In contrast, an estimate based from the biomass balance equation using measured carbon stocks and mortality was only 16.4 years. It is likely that the discrepancy of this third estimate with the first two estimates is due to CWD stocks measured in the inventory being too low. Coarse woody debris in natural forest is extremely difficult to measure accurately. We conclude that the methods used in the N.Z. natural forest inventory substantially underestimate CWD carbon stocks. Actual stocks of CWD are likely to average about 30 t C ha -1 compared with the measured value of 20 t C ha -1 . It is likely that the underestimate of CWD by the inventory is due to multiple factors. In particular, the measurement protocols exclude wood that is heavily decayed, bark and sapwood that has sloughed off stems, and dead wood buried in the forest floor. It is also probable that the density modifiers used in the analysis over-state the loss in density of decayed wood.

Comparison of measured and modelled change in coarse woody debris carbon stocks in New Zealand’s natural forest

Thermal refugia in cleared temperate Australian woodlands: Coarse woody debris moderate extreme surface soil temperatures

Hunter’s classification is the most adopted method for describing coarse woody debris (CWD) decay stages; however, wood chemical characterization is often lacking. This study explores the sugar profiles of living silver fir (Abies alba Mill.) trees and CWD samples in Hunter’s five-decay stages, evaluating the trends and variability in sugar content in CWD samples during decomposition. Wood cores from living trees and CWD samples from two different forest sites in Trentino (Italy) were analyzed after extraction using ion-exchange chromatography, equipped with pulsed amperometric and charge aerosol detectors (IC-PAD-CAD). Detailed monosaccharides, disaccharides, and sugar alcohol profiles of living wood and CWD samples were described. Cellobiose, arabinose, maltose, and trehalose showed similar trend in both the sites. Principal component analysis (PCA) highlighted the good capability of sugar profiles to well characterize and discriminate silver fir wood samples among Hunter’s categories of wood decomposition.

Research Highlights: Thinning and tree species alter the forest floor microclimate by modifying canopy cover, radiation, wind, and humidity. Thus, forest management can directly influence the edaphic mesofauna responsible for decomposing coarse woody debris (CWD). Background and Objectives: This research was carried out in the Southwestern Pyrenees Mountains (Northern Spain) and aimed to determine the influence of forest thinning and canopy type (pure Pinus sylvestris L. or a mix of P. sylvestris and Fagus sylvatica L.) on CWD colonization by edaphic fauna. Materials and Methods: CWD samples were collected belonging to intermediate and advanced decomposition stages, approximately 10 cm long and 5 cm in diameter. Using a design of three thinning intensities (0%, 20%, and 40% of basal area removed), with three replications per treatment (nine plots in total), four samples were taken per plot (two per canopy type) to reach 36 samples in total. Meso- and macrofauna were extracted from CWD samples with Berlese–Tullgren funnels, and individuals were counted and identified. Results: 19 taxonomic groups were recorded, the most abundant being the mesofauna (mites and Collembola). Mixed canopy type had a significant positive influence on richness, whereas advanced decay class had a positive significant influence on total abundance and richness. In addition, there were non-significant decreasing trends in richness and abundance with increasing thinning intensity. However, interactions among thinning intensity, canopy type, and decay class significantly affected mesofauna. Furthermore, some taxonomic groups showed differential responses to canopy type. CWD water content was positively correlated with total invertebrate abundance and some taxonomic groups. Our results suggest that stand composition has the potential to directly affect invertebrate communities in CWD, whereas stand density influence is indirect and mostly realized through changes in CWD moisture. As mesofauna is related to CWD decomposition rates, these effects should be accounted for when planning forest management transition from pure to mixed forests.

We investigated the mass dynamics of coarse woody debris (CWD) in an old-growth deciduous forest dominated by Quercus serrata, Carpinus laxiflora, and C. cordata in a 1 ha permanent plot of the Gwangneung Experiment Forest, Korea, from 2002 through 2010. CWD mass varied from 16.8 to 34.2 Mg ha−1, and the ratio of CWD mass to stand biomass varied from 0.06 to 0.13. The mean CWD mass input and loss rates were 4.81 Mg ha−1 yr−1 and 2.28 Mg ha−1 yr−1, respectively. A large heterogeneity of CWD mass, as represented by the spatial coefficient of variation (127.2%) and annual coefficient of variation (178.5%), might be inherent in the old-growth temperate forest, which consisted of large biomass trees. The decay rate constant, as estimated from the wood density change, was 0.049yr−1. However, the large variation of annual CWD mass input could cause the overestimation of decay rate constant (0.167 yr−1) as calculated from the ratio of CWD mass input to CWD mass. According to the CWD decay class classification, class II (72.8%) comprised the majority of CWD mass. The proportion of CWD mass to total CWD mass was 57.5% for Q. serrata, 25.0% for C. laxiflora, and 10.4% for C. cordata, respectively, and corresponded to the proportion of stem biomass to total stem biomass. These data support the stability of the current status in this old-growth deciduous forest as representing the climax stage. Due to the relatively short-term measurement of CWD mass compared with the whole life span of CWD, additional long-term studies with various approaches are required to enhance the knowledge of CWD mass dynamics in this forest.

This paper synthesizes data extracted from the literature and data collected in various studies by the author on the quantity, characteristics, and functional importance of coarse woody debris (CWD) in the old-growth forests of British Columbia (B.C.). There is little agreement in the literature about the minimum diameter of CWD or the number of decay classes recognized. In western North America, five decay classes are commonly used, but recent studies suggest fewer decay classes are preferable. Comparisons among decay classes and biogeoclimatic zones and subzones in B.C. reveal that quantities and volumes are greatest (up to approximately 60 kg/m2 and approximately 1800 m3/ha, respectively), and CWD persists the longest (sometimes in excess of 1000 years) in the Coastal Western Hemlock (CWH) biogeoclimatic zone. The quantity and ground cover of CWD increase with forest productivity. Persistence of CWD has varied from less than 100 to over 800 years in two coastal (CWH and Mountain Hemlock (MH)) and three interior (Interior Douglas-fir (IDF), Interior Cedar–Hemlock (ICH), and Engelmann Spruce – Subalpine Fir (ESSF)) biogeoclimatic zones. Trends in CWD quantity with forest age in managed coastal B.C. forests suggest a U-shaped curve, with greater quantities occurring in recent cutovers than in old-growth forests, and lowest quantities occurring in middle-aged forests. This may be the normal trend in CWD with forest age, with departures from this trend resulting from disturbance- or environment-specific factors. Relatively large amounts of data exist on the characteristics of CWD in the CWH, IDF, ICH, ESSF, and Boreal White and Black Spruce (BWBS) biogeoclimatic zones, but such data for the Coastal Douglas-fir, Sub-Boreal Pine–Spruce, Sub-Boreal Spruce (SBS), and Spruce–Willow–Birch biogeoclimatic zones appear relatively sparse. There have been few studies of the functional role of CWD in B.C. forests, but those studies that have been completed indicate that CWD is an important habitat component for some plant and animal species. A total of 169 plant species, including >95% of all lichens and liverworts, were found to grow on CWD in old-growth forests in the CWH, MH, IDF, ICH, and ESSF biogeoclimatic zones. One third of these species were restricted to CWD. Studies in several biogeoclimatic zones have found that CWD provided preferred habitat for and was associated with higher populations of some small animal species, such as shrews, some voles, and some salamanders, in old-growth forests, but the effects varied with species and biogeoclimatic zone. The nutrient cycling role of CWD is not yet well known, but it currently appears to be relatively insignificant in B.C. old-growth forests. Although it has been considered that CWD could increase mineral soil acidification and eluviation, no evidence for this was found in a study of the CWH, MH, IDF, ICH, ESSF, BWBS, and SBS biogeoclimatic zones. Future studies of the functional role of CWD should consider both scale (square metre vs. hectare) and temporal (changes in CWD with forest age) issues, as studies including these are sparse and both may be important. Key words: biogeoclimatic zones, British Columbia, coarse woody debris, old-growth forests.

Although coarse woody debris (CWD) is important for soil functioning, the mechanism which affects soil properties beneath CWD are unclear. Here, initial changes in microbial and soil properties were studied using homogenous CWD samples in eight Korean red pine (Pinus densiflora Sieb. et Zucc.) forests. For each forest, CWD samples (diameter: 11.1 ± 0.1 cm; length: 10.2 ± 0.0 cm) from similarly aged Korean red pine trees were laid on the mineral soil surface from May to June, 2016, and soils were sampled at points beneath CWD and at a distance of 1 m from the CWD after 1 year. Soils beneath the CWD had higher moisture but lower inorganic nitrogen (N) and a higher microbial biomass C (carbon)/N ratio than those sampled 1 m from the CWD. No differences in total C and N, labile C, pH, and C substrate utilization between the soils were significant. The difference in inorganic N between the soils decreased with increasing CWD decomposition, whereas that for microbial biomass fraction in total C and N increased correspondingly. Our results showed that soil microbial affinity for retaining N might become higher than that for retaining C under the presence of CWD, which possibly alters N availability and generates a spatial heterogeneity in forest soils.

We examined the species richness and host utilization patterns of wood-inhabiting aphyllophoraceous fungi (polypores and related fungi) in an old-growth beech and oak forest in a cool, temperate area of Japan. Coarse woody debris (CWD) ≥ 20 cm diam within a 6 ha plot was surveyed in Sep 2002. Tree genus, diameter, decay class and tree part of CWD samples were recorded. Fruiting bodies of aphyllophoraceous fungi that arose from the CWD were surveyed three times and identified to species. In total 256 CWD samples from 12 tree genera were surveyed with Quercus being the most frequent followed by Castanea and Fagus. From 196 CWD samples we recorded 436 wood-inhabiting fungi belonging to 63 species. Fifteen fungal species had at least 10 records, with Hymenochaete rubiginosa, Daedalea dickinsii, Xylobolus frustulatus, Rigidoporus cinereus and the small form of Fomes fomentarius being the most frequent. The number of fungal species that appeared on Fagus was significantly larger than that on Castanea, when the number of fruiting bodies collected was at least 50. The occurrences of the 15 dominant fungal species, except Trametes versicolor, were related to traits of the CWD. Tree genus was a predictor variable that affected the appearance of 11 of the 15 species of wood-inhabiting fungi. Only the tree part was selected for the models of Rigidoporus eminens, Schizopora flavipora and Stereum ostrea. Our results suggest that tree genus and tree part are important factors determining fungal community structure because these were selected as complementary predictor variables. Both oak and beech appear to be the most important tree genera for maintaining wood-inhabiting fungal species richness because the fungal flora formed on oak CWD is nearly complementary to those on chestnut, with low fungal species richness.

The role of coarse woody debris (CWD) in the global carbon (C) cycle is growing under increasing tree mortality driven by climate variability and disturbances. Quantifying C in CWD critically depends on accurate estimates of CWD density and C concentration in CWD. This study considered the main decomposition pathways (the proportions of CWD decomposed by fungi vs. invertebrates; fungal decay types) and the relationships between decay class, wood density, moisture, and C content of CWD in old‐growth mixed monsoon montane tropical forests in Vietnam based on the inventory of 359 CWD pieces. The bulk density of wood of the 1st, 2nd, 3rd, 4th, and 5th decay classes averaged 0.56, 0.49, 0.37, 0.28, and 0.15 g cm−3, respectively. The density reduction across decay classes did not significantly differ for snags, stumps, branches, leaning, and lying logs. Wood density was negatively related to wood moisture. The wood mass loss averaged 0%, 20%, 37%, 54%, and 74% in the CWD of the 1st, 2nd, 3rd, 4th, and 5th decay classes, respectively. Signs of invertebrate activity were recorded on 3% of CWD pieces. Among CWD samples with identifiable decay type, 88% were decomposed by white‐rot, 8% were decomposed as brown‐rot, and 4% of pieces contained both white‐ and brown‐rot patches. The mean C concentration in wood was 46.5%. It did not change with decay class. Our research provides an empirical basis for future inventories of CWD carbon stocks in Asian montane tropical mixed forests.

Fallen coarse woody debris (CWD) is critical to forest biodiversity and function. Few studies model factors that influence CWD availability, although such investigations are critically needed to inform sustainable forest management. We assess benchmark levels of CWD in unharvested native forests and those harvested for timber, across a range of forests in north‐east New South Wales, Australia. We found timber‐harvesting was the dominant driver of CWD, with almost double the count (pieces ha−1) and volume (m3 ha−1) of total CWD in selectively harvested than unharvested sites. This pattern was consistent across wet and dry forest types. Harvested sites had greater counts of hollow‐bearing logs, and greater volumes of small and medium‐sized CWD (15–50 cm diameter) than unharvested sites. There was no effect of harvesting on the volume of large CWD (>51 cm diameter). Total volumes of CWD (>15 cm diameter) varied from 114 to 166 m3 ha−1. We found few differences in CWD counts and volumes between forest types, with grassy woodlands and forests containing less CWD than other dry and shrubby forest types, reflecting lower potential input rates. The CWD levels recorded here are similar to those recorded in dry and wet sclerophyll forests elsewhere in Australia and are typical of global estimates for ‘old growth’ forests. Using general linear models we captured up to 57% of the variation in CWD across sites, and found that timber harvesting, topography and the numbers of standing hollow‐bearing and dead trees were significant predictors of CWD. Values for unharvested forest provide a benchmark that could be used to inform retention guidelines for CWD in managed forests in this region. Further assessment of the effect of repeat timber harvesting is needed to fully understand its impact on CWD dynamics, especially if forest residues resulting from timber harvesting are removed from native forests for bioenergy production.

Woody debris in treefall gaps shelters palatable plant species from deer browsing, in an old-growth temperate forest

Restoration of the pool of coarse woody debris after disturbances is one of the mechanisms for maintaining the stability of forest biogeocenoses. The studies of coarse woody debris have been carried out in the “Vepssky Forest” Reserve in the Leningrad Region on 8 sample plots established in primary forests (4 sample plots) and in secondary forests of the 1st generation after logging in 1973–1974 (4 sample plots), where the composition and structure of the stand, as well as the site conditions have been identical to those in primary forests. The coarse woody debris has been inventoried on transects. The stocks of coarse woody debris in primary stands have ranged from 104 to 233 m3 ha–1. Windfall and leaning trees have prevailed. The proportion of deadwood in both primary and secondary forests has been low. Clear cutting has significantly changed not only the stock of coarse woody debris, but also its distribution by decay classes and substrate categories. The stocks of coarse woody debris in secondary forests have ranged from 8 to 40 m3 ha–1, and have been mainly represented by stumps. The coarse woody debris of the 4th and 5th decay classes has almost been absent in primary forests, while in secondary forests the proportion of highly decomposed wood remaining after cutting has been about 50 %. The ratio of coarse woody debris and growing stocks has been on average 1:1 and 1:5 in the biogeocenoses of primary and secondary forests, respectively. The annual carbon balance of coarse woody debris (the difference in fluxes due to the loss of growing forest and the decay of coarse woody debris) has ranged from 0,40 to 2,80 t C ha–1 year–1, averaging 1,75 and 0,63 t C ha–1 year–1 in secondary and primary forests, respectively. The positive annual carbon balance in the coarse woody debris is due to the predominance of the rate of the loss of growing forest over the rate of the decay of coarse woody debris in primary forests as a result of wind disturbances and as a result of self-thinning of the stand in secondary forests.

In most temperate forest ecosystems, tree mortality over time generates downed logs that accumulate as coarse woody debris (CWD) on the forest floor. These downed logs and trunks have important recognized ecosystem functions including habitat for different organisms and long-term organic C storage. Due to its recalcitrant chemical composition and slow decomposition, CWD can also have direct effects on ecosystem carbon and nutrient turnover. CWD could also cause changes indirectly through the physical and chemical alterations that it generates, although it is not well-understood how important these indirect effects could be for ecosystem processes and soil biogeochemistry. We hypothesized that in an old-growth mature forest, CWD affects carbon and nutrient cycles through its “proximity effects”, meaning that the forest floor near CWD would have altered soil biotic activity due to the environmental and biogeochemical effects of the presence of CWD. We conducted our study in an old-growth southern beech temperate forest in Patagonia, Argentina, where we estimated and classified the distribution and mass, nutrient pools and decay stage of CWD on the forest floor, and evaluated its impact on litter decomposition, soil mites and soil enzymatic activity of carbon and phosphorus-degrading enzymes. We demonstrate here that CWD in this ecosystem represents an important organic carbon reservoir (85 Mg ha−1) and nitrogen pool (0.42 Mg ha−1), similar in magnitude to other old-growth forests of the Northern Hemisphere. In addition, we found significant proximity effects of CWD, with increased C-degrading soil enzyme activity, decreased mite abundance, and more rapid litter decomposition beneath highly decayed CWD. Considered at the ecosystem scale in this forest, the removal of CWD could cause a decrease of 6% in soil enzyme activity, particularly in the summer dry season, and nearly 15% in annual litter decomposition. We conclude that beyond the established importance of CWD as a long-term carbon reservoir and habitat, CWD contributes functionally to the forest floor by influencing the spatial heterogeneity of microbial activity and carbon and nutrient turnover. These proximity effects demonstrate the importance of maintenance of this ecosystem component and should be taken into consideration for management decisions pertaining to carbon sequestration and functional diversity in natural forest ecosystems.