Simulation and Analysis of the Effect of a Spruce Budworm Outbreak on Carbon Dynamics in Boreal Forests of Quebec

Ecosystems - In a climate change context, the sequestration of atmospheric carbon (C) in forests is key for achieving emission targets. It is thus critical to understand how large-scale...     

Modeling Spruce Budworm Population Revisited: Impact of Physiological Structure on Outbreak Control

Understanding the dynamics of spruce budworm population is very important for the protection of spruce and balsam fir trees of North American forests, and a full understanding of the dynamics requires careful consideration of the individual physiological structures that is essential for outbreak control. A model as a delay differential equation is derived from structured population system, and is validated by comparing simulation results with real data from the Green River area of New Brunswick (Canada) and with the periodic outbreaks widely observed. Analysis of the equilibrium stability and examination of the amplitudes and frequencies of periodic oscillations are conducted, and the effect of budworm control strategies such as mature population control, immature population control and predation by birds are assessed. Analysis and simulation results suggest that killing only budworm larvae might not be enough for the long-term control of the budworm population. Since the time required for development during the inactive stage (from egg to second instar caterpillar) causes periodic outbreak, a strategy of reducing budworms in the inactive stage, such as removing egg biomass, should also be implemented for successful control.     

Polyandry across Behavioral Classes in Female Spruce Budworm

Reproduction in spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), is characterized by sedentary oviposition in the natal forest patch early in life (gravid females are incapable of flight due to their heavy abdomen) followed by windborne migrations among females that have laid >50% of their eggs; the nature of migrations (facultative or obligatory) remains unknown. The incidence of polyandry (females with more than one spermatophore in their reproductive tract) was evaluated for several thousand field-collected female budworms (data professionally archived at Canadian Forest Service) using an array of sampling procedures that can be delineated along four behavioral classes: (1) resident females collected on host trees; (2) in-flight females captured while foraging in forest canopies; (3) migrant females captured above tree canopies; and (4) females collected postmortem on drop trays. Overall, polyandry increased over time (reflecting the ageing of local populations) and with female body size (likely due to male mating preference for large, most fecund females). Polyandry did not increase postmortem realized fecundity of females. The level of polyandry was lowest among migrants, intermediate for in-flight and resident females, and highest for naturally dead females. Sedentary reproduction among young female budworms may limit gene flow and increase the risk of inbreeding; a low level of polyandry among migrants may thus be due to females not only seeking novel habitats for their progeny but also mating partners that are not closely related genetically.     

Modelling Spatial Interactions Among Fire,Spruce Budworm,and Logging in the Boreal Forest

In the boreal forest, fire, insects, and logging all affect spatial patterns in forest age and species composition. In turn, spatial legacies in age and composition can facilitate or constrain further disturbances and have important consequences for forest spatial structure and sustainability. However, the complex three-way interactions among fire, insects, and logging and their combined effects on forest spatial structure have seldom been investigated. We used a spatially explicit landscape simulation model to examine these interactions. Specifically, we investigated how the amount and the spatial scale of logging (cutblock size) in combination with succession, fire, and spruce budworm outbreaks affect area burned and area defoliated. Simulations included 30 replicates of 300 years for each of 19 different disturbance scenarios. More disturbances increased both the fragmentation and the proportion of coniferous species and imposed additional constraints on the extent of each disturbance. We also found that harvesting legacies affect fire and budworm differently due to differences in forest types consumed by each disturbance. Contrary to expectation, budworm defoliation did not affect area burned at the temporal scales studied and neither amount of logging nor cutblock size influenced defoliation extent. Logging increased fire size through conversion of more of the landscape to early seral, highly flammable forest types. Although logging increased the amount of budworm host species, spruce budworm caused mortality was reduced due to reductions in forest age. In general, we found that spatial legacies do not influence all disturbances equally and the duration of a spatial legacy is limited when multiple disturbances are present. Further information on post-disturbance succession is still needed to refine our understanding of long-term disturbance interactions.     

Patterns of Carbon, Nitrogen and Phosphorus Dynamics in Decomposing Foliar Litter in Canadian Forests

We examined the patterns of nitrogen (N) and phosphorus (P) gain, retention or loss in ten foliar tissues in a litterbag experiment over 6 years at 18 upland forest sites in Canada, ranging from subarctic to cool temperate. N was usually retained in the decomposing litter until about 50% of the original C remained. The peak N content in the litter was observed at between 72 and 99% of the original C remaining, with C:N mass quotients between 37 and 71 (mean 55). The rate of N release from the litters was not related to the original N concentration, which may be associated with the generally narrow range (0.59–1.28% N) in the litters. P was immediately lost from all litters, except beech leaves, with critical litter C:P mass quotients for P release being in the range 700–900. The rate of P loss was inversely correlated with the original litter P concentration, which ranged from 0.02 to 0.13%. The soil underlying the litterbags influenced the pattern of N and P dynamics in the litters; there were weak correlations between the N and P remaining at 60% C remaining in the litters and the C:N and C:P quotients of the surface layer of the soil. There was a trend for higher N and P retention in the litter at sites with lower soil C:N and N:P quotients, respectively. Although there was a large variation in C:N, C:P and N:P quotients in the original litters (29–83, 369–2122 and 5–26, respectively), and some variation in the retention or loss of N and P in the early stages of decomposition, litters converged on C:N, C:P and N:P quotients of 30, 450 and 16, when the C remaining fell below 30%. These quotients are similar to that found in the surface organic matter of these ecosystems.     

Residence Times and Decay Rates of Downed Woody Debris Biomass/Carbon in Eastern US Forests

A key component in describing forest carbon (C) dynamics is the change in downed dead wood biomass through time. Specifically, there is a dearth of information regarding the residence time of downed woody debris (DWD), which may be reflected in the diversity of wood (for example, species, size, and stage of decay) and site attributes (for example, climate) across the study region of eastern US forests. The empirical assessment of DWD rate of decay and residence time is complicated by the decay process itself, as decomposing logs undergo not only a reduction in wood density over time but also reductions in biomass, shape, and size. Using DWD repeated measurements coupled with models to estimate durations in various stages of decay, estimates of DWD half-life (T _HALF), residence time (T _RES), and decay rate (k constants) were developed for 36 tree species common to eastern US forests. Results indicate that estimates for T _HALF averaged 18 and 10 years for conifers and hardwoods, respectively. Species that exhibited shorter T _HALF tended to display a shorter T _RES and larger k constants. Averages of T _RES ranged from 57 to 124 years for conifers and from 46 to 71 years for hardwoods, depending on the species and methodology for estimating DWD decomposition considered. Decay rate constants (k) increased with increasing temperature of climate zones and ranged from 0.024 to 0.040 for conifers and from 0.043 to 0.064 for hardwoods. These estimates could be incorporated into dynamic global vegetation models to elucidate the role of DWD in forest C dynamics.     

Tree Species Change in Nemoral Spruce Forests of the Central Forest Reserve

Biology Bulletin - Changes in the forest-forming species in the nemoral spruce forests of the Central Forest Reserve (Tver oblast, Russian Federation) are discussed. The characterization of...     

Control of Nitrous Oxide Emissions in European Beech,Norway Spruce and Scots Pine Forests

Elevated nitrogen deposition has increased tree growth, the storage of soil organic matter, and nitrate leaching in many European forests, but little is known about the effect of tree species and nitrogen deposition on nitrous oxide emission. Here we report soil N₂O emission from European beech, Scots pine and Norway spruce forests in two study areas of Germany with distinct climate, N deposition and soils. N₂O emissions and throughfall input of nitrate and ammonium were measured biweekly during growing season and monthly during dormant season over a 28 months period. Annual N₂O emission rates ranged between 0.4 and 1.3 kg N ha^?1 year^?1 among the stands and were higher in 1998 than in 1999 due to higher precipitation during the growing season of 1998. A 2-way-ANOVA revealed that N₂O fluxes were significantly higher (p<0.001) at Solling than at Unterlüß while tree species had no effect on N₂O emissions. Soil texture and the amount of throughfall explained together 94% of the variance among the stands, indicating that increasing portions of silt and clay may promote the formation of N₂O in wet forest soils. Moreover, cumulative N₂O fluxes were significantly correlated (r² = 0.60, p<0.001) with cumulative NO ₃ ^? fluxes at 10 cm depth as an indicator of N saturation, however, the slope of the regression curve indicates a rather weak effect of NO ₃ ^? fluxes on N₂O emissions. N input by throughfall was not correlated with N₂O emissions and only 1.6–3.2% of N input was released as N₂O to the atmosphere. Our results suggest that elevated N inputs have little effect on N₂O emissions in beech, spruce and pine forests.     

Impact of Future Climate on Radial Growth of Four Major Boreal Tree Species in the Eastern Canadian Boreal Forest

Immediate phenotypic variation and the lagged effect of evolutionary adaptation to climate change appear to be two key processes in tree responses to climate warming. This study examines these components in two types of growth models for predicting the 2010–2099 diameter growth change of four major boreal species Betula papyrifera, Pinus banksiana, Picea mariana, and Populus tremuloides along a broad latitudinal gradient in eastern Canada under future climate projections. Climate-growth response models for 34 stands over nine latitudes were calibrated and cross-validated. An adaptive response model (A-model), in which the climate-growth relationship varies over time, and a fixed response model (F-model), in which the relationship is constant over time, were constructed to predict future growth. For the former, we examined how future growth of stands in northern latitudes could be forecasted using growth-climate equations derived from stands currently growing in southern latitudes assuming that current climate in southern locations provide an analogue for future conditions in the north. For the latter, we tested if future growth of stands would be maximally predicted using the growth-climate equation obtained from the given local stand assuming a lagged response to climate due to genetic constraints. Both models predicted a large growth increase in northern stands due to more benign temperatures, whereas there was a minimal growth change in southern stands due to potentially warm-temperature induced drought-stress. The A-model demonstrates a changing environment whereas the F-model highlights a constant growth response to future warming. As time elapses we can predict a gradual transition between a response to climate associated with the current conditions (F-model) to a more adapted response to future climate (A-model). Our modeling approach provides a template to predict tree growth response to climate warming at mid-high latitudes of the Northern Hemisphere.     

Broadleaf Litter Controls Feather Moss Growth in Black Spruce and Birch Forests of Interior Alaska

Ecosystems - Plant–soil feedbacks can maintain or reinforce alternative states within ecological systems. In Alaskan boreal forests, changes in fire characteristics have stimulated the...     

Modelling Growth-Competition Relationships in Trembling Aspen and White Spruce Mixed Boreal Forests of Western Canada

We examined the effect of competition on stem growth of Picea glauca and Populus tremuloides in boreal mixedwood stands during the stem exclusion stage. We combined traditional approaches of collecting competition data with dendrochronology to provide retrospective measurements of stem diameter growth. Several competition indices including stand basal area (BA), the sum of stem diameter at breast height (SDBH), and density (N) for the broadleaf and coniferous species, as well as similar indices considering only trees with diameters greater than each subject (BAGR, SDBHGR, and NGR), were evaluated. We used a nonlinear mixed model to characterize the basal area increment over the past 5, 10, 15, 20, 25, 30, and 35 years as a function of growth of nearby dominant trees, the size of the subject trees, deciduous and coniferous competition indices, and ecoregions. SDBHGR and BAGR were better predictors for spruce, and SDBHGR and NGR were better for aspen, respectively, than other indices. Results showed strongest correlations with long-term stem growth, as the best models integrated growth for 10–25 years for aspen and ≥25 for spruce. Our model demonstrated a remarkable capability (adjusted R²>0.67) to represent this complex variation in growth as a function of site, size and competition.     

Classification of Morphogenetic Types of Mossy Litter in Paludine Spruce Forests Based on Humus Content

We have classified morphogenetic types of mossy litter by a multivariate statistical analysis of a fractional group of organic-matter composition. Three clusters of mossy litters—peats, peaty, and high-ash peaty— are recognized. This results in 94% prediction. Indicators contribute to discriminants in the following descending order: C: N > cellulose > HA-2 > HA-3 > FA-3 > hemicellulose = FA-1 = HA-1. According to canonical analysis, there were two significant roots in cluster determination. The first segregates mainly the peat cluster from two others. The share of canonical function of the root one is 58% of possible discrimination, mainly due to the weight of cellulose and C: N. Canonical function 2, describing 42% of the explained dispersion, discriminates the peaty cluster from the others due to the dominant contribution of FA-1 and FA-3. The classification function for the recognition of new objects was calculated and evaluated. The humus content of various types and clusters of mossy litters was examined. Morphogenetic classification follows the transformation of forest litter in the course of litter formation (continuum phase), quantitative biochemical discrimination is a discrete phase of this process.     

Review of Carbon Fixation in Bamboo Forests in China

Bamboo is widespread in the subtropics and tropics of Asia, Africa, and Latin America. The total area of bamboo forests of various species is 22.0?×?10⁶ ha, accounting for about 1.0% of the total global area of forest. Although the total forest areas in many countries have decreased drastically, bamboo forests have increased at a rate of 3% annually. China has the richest resources of bamboo in the world, with over 500 species in 39 genera. Carbon storage of vegetation, soils, and litter in bamboo forest system in China was 0.2511?×?10¹⁵, 0.8516?×?10¹⁵, and 0.0361?×?10¹⁵ g C, respectively, giving a total of 1.1388?×?10¹⁵ g C. This paper reviews carbon storage of vegetation, soils, and litter in bamboo forest system and compares the carbon fixation abilities of bamboo forest ecosystems with those of other tree species in subtropical China.     

Cry9Ca1 Toxin, a Bacillus thuringiensis Insecticidal Crystal Protein with High Activity against the Spruce Budworm (Choristoneura fumiferana)

The Cry9Ca1 toxin from Bacillus thuringiensis was significantly more toxic to spruce budworm (Choristoneura fumiferana) than the Cry1Ab6, Cry1Ba1, Cry1Ca2, Cry1Da1, Cry1Ea1, and Cry1Fa2 toxins. It displayed high activity against silkworm (Bombyx mori) but was not toxic to black army cutworm (Actebia fennica) or gypsy moth (Lymantria dispar). The Cry9Ca1 is the most effective spruce budworm toxin known to date and may offer promise for control and resistance management of that species.     

Soil Respiration and Belowground Carbon Allocation in Mangrove Forests

Mangrove forests cover large areas of tropical and subtropical coastlines. They provide a wide range of ecosystem services that includes carbon storage in above- and below ground biomass and in soils. Carbon dioxide (CO₂) emissions from soil, or soil respiration is important in the global carbon budget and is sensitive to increasing global temperature. To understand the magnitude of mangrove soil respiration and the influence of forest structure and temperature on the variation in mangrove soil respiration I assessed soil respiration at eleven mangrove sites, ranging from latitude 27°N to 37°S. Mangrove soil respiration was similar to those observed for terrestrial forest soils. Soil respiration was correlated with leaf area index (LAI) and aboveground net primary production (litterfall), which should aid scaling up to regional and global estimates of soil respiration. Using a carbon balance model, total belowground carbon allocation (TBCA) per unit litterfall was similar in tall mangrove forests as observed in terrestrial forests, but in scrub mangrove forests TBCA per unit litter fall was greater than in terrestrial forests, suggesting mangroves allocate a large proportion of their fixed carbon below ground under unfavorable environmental conditions. The response of soil respiration to soil temperature was not a linear function of temperature. At temperatures below 26°C Q10 of mangrove soil respiration was 2.6, similar to that reported for terrestrial forest soils. However in scrub forests soil respiration declined with increasing soil temperature, largely because of reduced canopy cover and enhanced activity of photosynthetic benthic microbial communities.     

Significance of Over-Mature and Decaying Trees for Carbon Stocks in a Central European Natural Spruce Forest

Old-growth forests are important stores for carbon as they may accumulate C for centuries. The alteration of biomass and soil carbon pools across the development stages of a forest dynamics cycle has rarely been quantified. We studied the above- and belowground C stocks in the five forest development stages (regeneration to decay stage) of a montane spruce (Picea abies) forest of the northern German Harz Mountains, one of Central Europe’s few forests where the natural forest dynamics have not been disturbed by man for several centuries. The over-mature and decay stages had the largest total (up to 480 Mg C ha^?1) and aboveground biomass carbon pools (200 Mg C ha^?1) with biomass C stored in dead wood in the decay stage. The soil C pool (220–275 Mg C ha^?1, 0–60 cm) was two to three times larger than in temperate lowland spruce forests and remained invariant across the forest dynamics cycle. On the landscape level, taking into account the frequency of the five forest development stages, the total carbon pool was approximately 420 Mg C ha^?1. The results evidence the high significance of over-mature and decaying stages of temperate mountain forests not only for conserving specialized forest organisms but also for their large carbon storage potential.     

Availability of Ectomycorrhizal Fungi to Black Spruce above the Present Treeline in Eastern Labrador

Ectomycorrhizal fungi (ECMF) are an important biotic factor in the survival of conifer seedlings under stressful conditions and therefore have the potential to facilitate conifer establishment into alpine and tundra habitats. In order to assess patterns of ectomycorrhizal availability and community structure above treeline, we conducted soil bioassays in which Picea mariana (black spruce) seedlings were grown in field-collected soils under controlled conditions. Soils were collected from distinct alpine habitats, each dominated by a different ectomycorrhizal host shrub: Betula glandulosa, Arctostaphylos alpina or Salix herbacaea. Within each habitat, half of the soils collected contained roots of ectomycorrhizal shrubs (host⁺) and the other half were free of host plants (host⁻). Forest and glacial moraine soils were also included for comparison. Fungi forming ectomycorrhizae during the bioassays were identified by DNA sequencing. Our results indicate that ECMF capable of colonizing black spruce are widespread above the current tree line in Eastern Labrador and that the level of available inoculum has a significant influence on the growth of seedlings under controlled conditions. Many of the host⁻ soils possessed appreciable levels of ectomycorrhizal inoculum, likely in the form of spore banks. Inoculum levels in these soils may be influenced by spore production from neighboring soils where ectomycorrhizal shrubs are present. Under predicted temperature increases, ectomycorrhizal inoculum in soils with host shrubs as well as in nearby soils without host shrubs have the potential to facilitate conifer establishment above the present tree line.     

Convergence in Arthropod Assemblages with Various Restoration Approaches for Canadian Deciduous Forests

Silvicultural practices are traditionally aimed at increasing forest profits; however, recent approaches to forest conservation have broadened to include nature-based silviculture for regenerating forests. In southern Ontario (Canada), originally dominated by deciduous forests, conifer plantations were established on abandoned agricultural sites. Currently, there is an increasing interest to convert these conifer stands to a state that mimics the original deciduous forest. We investigated arthropod abundance, species richness of carabid beetles, and abundance of arthropod assemblages (trophic and prey groups) under five silvicultural treatments conducted to regenerate deciduous forests (the natural forest type) from the old conifer plantations. The treatments included: (1) uniform canopy removal; (2) uniform canopy removal and understory removal; (3) group canopy removal; (4) group canopy removal and understory removal; and (5) untreated control plots (relatively pure red pine). Insects were sampled annually using sweepnets and pitfall traps. Results revealed treatment effects on the abundance of Coleoptera, Heteroptera, herbivores, and small arthropods (<3 mm) caught in sweepnet samples, where plots subjected to group shelterwood removal and understory removal supported higher abundances than the control plots. There was no treatment effect on the abundance of other arthropod groups or on the species richness and abundance of carabid beetles. The silvicultural treatments used to encourage natural regeneration did not seem to affect arthropod food availability for insectivorous vertebrates. Thus, the type of silvicultural strategy used to convert pine plantations to a stage that mimics the natural deciduous forests had little overall impact on arthropods.