Predicting the composition of Canadian southern boreal forest in different fire cycles

Abstract. Post-fire succession was reconstructed for a sector located in the southern part of the Québec boreal forest. Forest composition for different periods since fire was evaluated using a stand initiation map together with ecological maps representing both site conditions and stand types. Nine fires covering at least 100 ha and representing a chronosequence of more than 230 yr were used. Although a relatively clear successional pattern from deciduous to coniferous composition relating to time-since-fire was observed, Pinus banksiana stands showed an erratic distribution not related to succession but possibly to the pre-fire stand composition. A comparison with forest cover maps produced after a recent spruce budworm outbreak, showed that succession toward coniferous dominance appeared to be interrupted by spruce budworm (Choristoneura fumiferana) outbreaks which, by killing Abies balsamea, lead to a mixed deciduous forest composition. A simple empirical model based on a negative exponential distribution of age classes was developed to evaluate how changes in the fire cycle would affect the composition of the forest mosaic. The transition between deciduous dominance and coniferous dominance occurs in a fire cycle > 200 yr. Although pure deciduous stands tend to disappear during long fire cycles, the proportion of mixed stands remains relatively constant. Prediction of the forest composition for longer fire cycles is complicated by the interaction between post-fire composition and stand vulnerability to spruce budworm outbreaks.     

Spruce budworm impact, abundance and parasitism rate in a patchy landscape

The hypothesis that vegetational diversity may lessen the impact of forest insect pests by favoring natural enemies is appealing to those who seek ecologically sound solutions to pest problems. We investigated the effect of forest diversity on the impact of the spruce budworm Choristoneurafumiferana following the last outbreak, as well as the budworm's current abundance and parasitism rate, in the boreal forest of northwestern Québec. Mortality of balsam fir caused by the budworm was greater in extensive conifer stands than either in “habitat islands” of fir surrounded by deciduous forest or on true islands in the middle of a lake. Adult spruce budworm abundance, assessed by pheromone traps, did not differ significantly between the three types of sites. Larval and pupal parasitism rates were examined by transferring cohorts of laboratory-reared larvae and pupae to trees in the three site types and later collecting and rearing them. The tachinid Actiainterrupta, a parasitoid of fifth and sixth instar larvae, as well as the ichneumonid pupal parasitoids Itoplectesconquisitor, Ephialtesontario and Phaeogenesmaculicornis, caused higher mortality in the habitat islands than on true islands or in extensive stands. Exochusnigripalpistectulum, an ichneumonid that attacks the larvae and emerges from the pupae, caused greater mortality in the extensive stands of conifers. Received: 17 March 1997 / Accepted: 18 November 1997     

Forest composition, host-population density, and parasitism of spruce budworm Choristoneura fumiferana eggs by Trichogramma minutum

The frequency of egg parasitism in the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) by Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) was recorded in several host populations during the rising phase of an outbreak in the Gatineau Valley (Quebec, Canada) in 1999 and 2000. Some of the observed spruce budworm populations were subjected to experimental aerial applications of Bacillus thuringiensis var. kurstaki during the course of the study. Untreated spruce budworm population densities at the onset of the study were directly related to stand composition (the proportion of non‐host basal area) but unrelated to plant species diversity. Much of the variation in budworm egg mass parasitism rates was accounted for (77.1% of total variance) by three variables: year, spruce budworm egg population density, and proportion of non‐host basal area in the stand. The exploitation of eggs within parasitised egg masses was related to year, B. thuringiensis treatment, egg mass size, and the proportion of egg masses parasitised in the host population (76.9% of variance). The yearly variation in parasitism rate was considerable, and was consistent with climatic conditions, namely heat accumulation and precipitation during the spruce budworm oviposition period. Parasitism by T. minutum on eggs of C. fumiferana was recorded in 16 additional sites in 2000, to confirm the general form of the inverse density dependence of parasitism. Observations recorded by Neilson in the Green River area of New Brunswick (Canada) between 1948 and 1957 are discussed in the context of the present analysis. These results indicate that parasitism by T. minutum may be more predictable than previously believed and may be an important and consistent source of mortality in the several years preceding an outbreak of C. fumiferana, especially in mixed‐wood stands and in warmer, drier portions of the spruce budworm's range.     

Effects of fire and spruce beetle outbreak legacies on the disturbance regime of a subalpine forest in Colorado

Aim There is increasing research attention being given to the role of interactions among natural disturbances in ecosystem processes. We studied the interactions between fire and spruce beetle (Dendroctonus rufipennis Kirkby) disturbances in a Colorado subalpine forest. The central questions of this research were: (1) How does fire history influence stand susceptibility to beetle outbreak? And conversely, (2) How does prior occurrence of a beetle outbreak influence stand susceptibility to subsequent fire? Methods We reconstructed the spatial disturbance history in a c. 4600 ha area by first identifying distinct patches in the landscape on aerial photographs. Then, in the field we determined the disturbance history of each patch by dating stand origin, fire scars, dates of mortality of dead trees, and releases on remnant trees. A geographical information system (GIS) was used to overlay disturbance by fire and spruce beetle. Results and main conclusions The majority of stands in the study area arose following large, infrequent, severe fires occurring in c. 1700, 1796 and 1880. The study area was also affected by a severe spruce beetle outbreak in the 1940s and a subsequent low‐severity fire. Stands that originated following stand‐replacing fire in the late nineteenth century were less affected by the beetle outbreak than older stands. Following the beetle outbreak, stands less affected by the outbreak were more affected by low‐severity fire than stands more severely affected by the outbreak. The reduced susceptibility to low‐severity fire possibly resulted from increased moisture on the forest floor following beetle outbreak. The landscape mosaic of this subalpine forest was strongly influenced by the interactions between fire and insect disturbances.     

Not just for the birds: Spiders as natural enemies of spruce budworm (Choristoneura fumiferana,Clem.)

(1) The eastern spruce budworm (Choristoneura fumiferana, Clem.) is a native irruptive forest pest that defoliates spruce-fir forests throughout North America's boreal zone. (2) Past studies suggest that successful spruce budworm population control requires high natural mortality from a variety of sources, including predators, especially from parasitoids and birds. While well represented in many different ecosystems, the role of generalist predatory spiders in these boreal systems remains largely unstudied. (3) To determine the identity and percentage of spiders preying upon spruce budworm, we hand-collected spiders from balsam fir (Abies balsamea) in stands with relatively high spruce budworm densities from forests in insular Newfoundland and Labrador, Canada. (4) Using a spruce budworm-specific TaqMan real-time PCR assay, we successfully amplified spruce budworm DNA in 32% of collected spiders. After spider molecular barcoding, we found the web builders Grammonota angusta Dondale, Pityohyphantes (aff. subarcticus), Dictyna brevitarsa Emerton and Estrandia grandaeva (Keyserling) represented 58% of the spiders feeding on spruce budworm, and the wandering hunter Philodromus rufus Dondale represented 11.8%. (5) Our molecular approach was an effective means with which to identify recently consumed prey and natural enemies in this boreal system.     

Succession in the southern part of the Canadian boreal forest

Forest succession following fire in a forest mosaic of northwestern Quebec has been studied in order to: (1) describe the successional pathways using communities of different ages and (2) evaluate convergence of successional pathways and possible effect of fire suppression on the establishment of steady-state communities. As a first step, ordination and classification techniques were used in order to remove changes in forest composition which are related to abiotic conditions. Then, ordinations based on tree diameter distributions were used to study shifts in species composition in relation to time since the last fire.Even under similar abiotic conditions, successional pathways are numerous. However, regardless of forest composition after fire, most stands show convergence toward dominance of Thuja occidentalis and Picea mariana on xeric sites and dominance of Abies balsamea and Thuja occidentalis on more mesic sites. Stable communities of >300 yr occur on xeric sites while on mesic sites directional succession still occurs after 224 yr. Nearly all species involved in succession are present in the first 50 yr following fire. Only Abies balsamea and Thuja occidentalis increase significantly in frequency during succession. Following initial establishment, successional processes can generally be explained by species longevity and shade tolerance. Early successional species may be abundant in the canopy for more than 200 yr while the rapid decrease of Picea glauca, a late successional species could be related to spruce budworm outbreaks. Considering the short fire rotation observed (about 150 yr), a steady-state forest is unlikely to occur under natural conditions, though it may be possible if fire is controlled.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Future Spruce Budworm Outbreak May Create a Carbon Source in Eastern Canadian Forests

Spruce budworm (Choristoneura fumiferana Clem.) is an important and recurrent disturbance throughout spruce (Picea sp.) and balsam fir (Abies balsamea L.) dominated forests of North America. Forest carbon (C) dynamics in these ecosystems are affected during insect outbreaks because millions of square kilometers of forest suffer growth loss and mortality. We tested the hypothesis that a spruce budworm outbreak similar to those in the past could switch a forest from a C sink to a source in the near future. We used a model of ecosystem C to integrate past spruce budworm impact sequences with current forest management data on 106,000 km² of forest in eastern Québec. Spruce budworm-caused mortality decreased stand-level merchantable C stocks by 11–90% and decreased ecosystem C stocks by 2–10% by the end of the simulation. For the first 13 years (2011–2024), adding spruce budworm significantly reduced ecosystem C stock change for the landscape from a sink (4.6 ± 2.7 g C m⁻² y⁻¹ in 2018) to a source (−16.8 ± 3.0 g C m⁻² y⁻¹ in 2018). This result was mostly due to reduced net primary production. The ecosystem stock change was reduced on average by 2 Tg C y⁻¹ for the entire simulated area. This study provides the first estimate that spruce budworm can significantly affect the C sink or source status of a large landscape. These results indicate that reducing spruce budworm impacts on timber may also provide an opportunity to mitigate a C source.     

Potential of Margosan-O, an azadirachtin-containing formulation from neem seed extract, as a control agent for spruce budworm, Choristoneura fumiferana

The effects on spruce budworm larvae, Choristoneura fumiferana (Clem.), produced by ingestion of Margosan-O, a commercially available neem seed extract formulation containing 0.3% azadirachtin, were investigated. Bioassays with the test material were conducted using various instars of spruce budworm larvae, with either artificial diet, cut branches of balsam fir, Abies balsamea (L.) Mill., or small growing balsam fir trees as substrates. The dose-response data on feeding reduction, developmental retardation, and mortality (LC50, LC95, and LD50) suggest that Margosan-O has promise as a control agent for spruce budworm in an integrated pest management program.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Macrocharcoal-Based Chronosequences Reveal Shifting Dominance of Conifer Boreal Forests Under Changing Fire Regime

Balsam fir (Abies balsamea) and black spruce (Picea mariana) forests are the main conifer forest types in the North American boreal zone. The coexistence of the two species as well as their respective canopy dominance in distinct stands raises questions about the long-term evolution from one forest type to the other in relation to environmental factors including climate and stand disturbance. We tested the hypothesis that repetitive fire events promote the succession of balsam fir forest to black spruce forest and vice versa. Postfire chronosequences of one black spruce (BSP) and one balsam fir (BFI) sites were reconstructed based on the botanical composition and ¹⁴C-dated soil macrocharcoals. The results support the hypothesis of a successional dynamics. The BSP site has been affected by fires for the last 7600 years, whereas the BFI site, after having been impacted by several fires during the first half of the Holocene, evolved in a fire-free environment for the last 4400 years. Periods of fire activity facilitated the dominance of black spruce forests. The cessation of fires around 4400 cal. years BP on BFI site marks the beginning of the transition from black spruce to balsam fir stands. This succession is a long process, due to the ability of black spruce to regenerate by layering in the absence of fire. The resulting balsam fir stands are ancient and precarious ecosystems, since fire generally leads to the return of black spruce. The increase in balsam fir to the detriment of black spruce in boreal forests is a response to a decrease in fire frequency.     

Variation in wood anatomical structure of Douglas-fir defoliated by the western spruce budworm: a case study in the coastal-transitional zone of British Columbia,Canada

Key message

An outbreak of the western spruce budworm temporarily modifies cellular wood anatomy of stem wood in natural and mature Douglas-fir stands impacting wood quality properties.

Abstract

Western spruce budworm (Choristoneura occidentalis Freeman) is a widespread and destructive defoliator of commercially important coniferous forests in western North America. In British Columbia, Canada, Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] is the most important and widely distributed host. Permanent sample plots were established at a number of locations in southern interior at the beginning of a severe western spruce budworm outbreak in the 1970s. Two of the sites were sampled in 2012 to determine whether modifications had occurred in the anatomical characteristics of stem wood formed during outbreak years. We determined that rings formed during the western spruce budworm 1976–1980 outbreak had a significantly lower proportion of latewood, reduced mean cell wall thickness and smaller radial cell diameters. While the cellular characteristics of the earlywood remained fairly constant, significant reductions in lumen area occurred in 1978 and 1979 at each site. Our study shows that western spruce budworm outbreaks not only reduce annual radial growth, but also temporarily modify cellular characteristics in latewood cells, which has implications for wood density and quality in Douglas-fir.     

The influence of natural disturbances on developmental patterns in Acadian mixedwood forests from 1946 to 2008

We used dendrochronological analyses to identify periods of natural disturbance and resulting growth responses in 32 mixed species stands in the Acadian forest. Stands represented five different development patterns, based upon 1946 softwood (SW) content (70–80%, termed SW versus 30–60%, termed mixedwood (MW)) and change in SW content from 1946 to 2006: SW-stable, SW-declining, MW-fluctuating, MW-stable, and MW-declining. Standardized growth chronologies were developed from 1163 increment cores sampled from balsam fir (Abies balsamea), red spruce (Picea rubens), yellow birch (Betula alleghaniensis), and sugar maple (Acer saccharum). Growth chronologies clearly identified three spruce budworm (Choristoneura fumiferana) outbreaks from 1914 to 1921, 1954 to 1961, and 1975 to 1984, and birch dieback from 1938 to 1948. Stand developmental patterns were caused by species characteristics and multiple interacting disturbances, resulting in mortality, growth reductions, releases, and establishment of new cohorts. Balsam fir was present in all stands, but its tendency to establish and release from advanced regeneration following budworm-caused mortality resulted in cyclical proportions of fir in the canopy. Red spruce was less vulnerable to spruce budworm and longer lived, allowing them to persist and better withstand disturbance. Periodic growth index values less than 0.9 for fir and spruce were correlated with mortality of softwoods caused by defoliation, which resulted in release and growth index values >1.1 for sugar maple. Our results demonstrated substantial variation in mixedwood development patterns over a 60-year period within a small (45 km) area.     

Effects of bud phenology and foliage chemistry of balsam fir and white spruce trees on the efficacy of Bacillus thuringiensis against the spruce budworm, Choristoneura fumiferana

Abstract 1 Efficacy of commercial formulations of Bacillus thuringiensis ssp. kurstaki (Btk) against spruce budworm Choristoneura fumiferana was investigated in mixed balsam fir‐white spruce stands. Btk treatments were scheduled to coincide with early flaring of balsam fir shoots, and later with flaring of white spruce shoots. Btk efficacy on the two host trees was compared and examined according to the foliar content of nutrients and allelochemicals and the insect developmental stage at the time of spray. 2 Larvae fed white spruce foliage were less vulnerable to Btk ingestion than larvae fed balsam fir foliage. Higher larval survival on white spruce, observed 10 days after spray, was related to higher foliage content in tannins and a lower N/tannins ratio, which might have induced inactivation of Btk toxins. 3 Larval mortality due to Btk did not depend on spruce budworm larval age. 4 Foliage protection of both host trees was similar in plots treated with Btk: larval mortality due to Btk treatment reduced insect grazing pressure on balsam fir trees; meanwhile, suitability of white spruce foliage seemed to decrease very rapidly, which induced high larval mortality among spruce budworm fed on white spruce trees. Nevertheless, following Btk sprays, 50% more foliage remained on white spruce than on balsam fir trees, because of the higher white spruce foliage production. 5 Both spray timings achieved similar protection of white spruce trees, but Btk treatments had to be applied as early as possible (i.e. during the flaring of balsam fir shoots to optimally protect balsam fir trees in mixed balsam fir‐white spruce stands).     

Potential Role of Spruce Budworm in Range-Wide Decline of Canada Warbler

ABSTRACT The Canada warbler (Wilsonia canadensis) is one of many common neotropical migrants whose populations are in decline across their range. Influences of habitat loss and degradation on breeding or wintering grounds have been postulated as possible causes, but few empirical data exist to support a specific cause. Based on previous studies linking abundances of Canada warbler and spruce budworm (Choristoneura fumiferana), we hypothesized that the Canada warbler may be influenced by a persistent decline in spruce budworm throughout the bird's breeding range, a hypothesis that has received little attention. This hypothesis makes 5 predictions: 1) budworm outbreaks and warbler detections should be spatially and temporally coincident; 2) the relationship between Canada warbler and spruce budworm outbreaks should be similar to relationships for other warblers known to be spruce budworm associates; 3) the relationship should be stronger than for warblers lacking an association with spruce budworm; 4) because temporal trends of both spruce budworm and Canada warblers have varied throughout Canadian provinces, declines in Canada warblers should be seen only in provinces where spruce budworm also declined; and 5) variation in Canada warbler abundance should reflect variation in supply of preferred habitat for the spruce budworm if habitat rather than budworm abundance is the key. Our analyses supported predictions 1–4, suggesting that Canada warbler may be even more closely associated with spruce budworm than are known associated species, a phenomenon noted in the literature but previously unexplained. Prediction 5 was not supported, because budworm habitat (area of mature and older balsam fir [Abies balsamea] and white spruce [Picea glauca]) remained constant in Ontario while warbler abundance declined. Although the correlative nature of these results precludes inference of a causal relationship between the declines of the Canada warbler and spruce budworm, we postulate that potential links may exist directly, where spruce budworm outbreaks provide elevated levels of insect prey items for breeding Canada warblers, or indirectly through changes in forest structure and composition following outbreaks. These results have implications when considering long-term trends in Canada warbler populations, because it may be impossible to alter population trends for species linked to the timing and magnitude of spruce budworm outbreaks.     

Insect defoliation modulates influence of climate on the growth of tree species in the boreal mixed forests of eastern Canada

Increasing air temperatures and changing precipitation patterns due to climate change can affect tree growth in boreal forests. Periodic insect outbreaks affect the growth trajectory of trees, making it difficult to quantify the climate signal in growth dynamics at scales longer than a year. We studied climate‐driven growth trends and the influence of spruce budworm (Choristoneura fumiferana Clem.) outbreaks on these trends by analyzing the basal area increment (BAI) of 2058 trees of Abies balsamea (L.) Mill., Picea glauca (Moench) Voss, Thuja occidentalis L., Populus tremuloides Michx., and Betula papyrifera Marsh, which co‐occurs in the boreal mixedwood forests of western Quebec. We used a generalized additive mixed model (GAMM) to analyze species‐specific trends in BAI dynamics from 1967 to 1991. The model relied on tree size, cambial age, degree of spruce budworm defoliation, and seasonal climatic variables. Overall, we observed a decreasing growth rate of the spruce budworm host species, A. balsamea and P. glauca between 1967 and 1991, and an increasing growth rate for the non‐host, P. tremuloides, B. papyrifera, and T. occidentalis. Our results suggest that insect outbreaks may offset growth increases resulting from a warmer climate. The observation warrants the inclusion of the spruce budworm defoliation into models predicting future forest productivity.     

A dendroecological reconstruction of western spruce budworm outbreaks (Choristoneura occidentalis) in the Front Range,Colorado, from 1720 to 1986

Radial increment cores from Douglas-fir (Pseudotsuga menziesii) and blue spruce (Picea pungens), defoliated by western spruce budworm (Choristoneura occidentalis), were analyzed by means of dendrochronological methods and compared with cores from undefoliated ponderosa pine (Pinus ponderosa) and lodgepole pine (Pinus contorta) growing on the same sites in the Front Range, Colorado. Extensive deforestation during the gold and silver booms in the second part of the nineteenth century led to dense and almost pure stands of shadetolerant budworm host species. By using the skeleton plot method, the number of trees with clear growth reductions is obtained, thus representing an exact record of forest insect attacks. The analysis of abrupt growth reductions revealed at least nine outbreaks of western spruce budworm between 1720 and 1986, the majority occurring in the nineteenth century. The outbreaks were graphically compared with periods of attack in New Mexico and Colorado which were detected by other scientists employing tree-ring measurement techniques. No increase in the frequency of severe outbreaks during the twentieth century was observed, yet there is some evidence that the most recent outbreak might be the most severe ever recorded. Open Douglas-fir stands on higher sites were more susceptible to heavy budworm attack than dense stands on lower sites. Blue spruce was less frequently and less severely attacked than Douglas-fir. The spatial pattern of historical outbreaks generally was very patchy.     

Interspecific variation in resistance of two host tree species to spruce budworm

Woody plants regularly sustain biomass losses to herbivorous insects. Consequently, they have developed various resistance mechanisms to cope with insect attack. However, these mechanisms of defense and how they are affected by resource availability are not well understood. The present study aimed at evaluating and comparing the natural resistance (antibiosis and tolerance) of balsam fir (Abies balsamea [L.] Mill.) and white spruce (Picea glauca [Moench) Voss] to spruce budworm, Choristoneura fumiferana (Clem.), and how drainage site quality as a component of resource availability affects the expression of resistance over time (6 years). Our results showed that there are differences in natural resistance between the two tree species to spruce budworm, but it was not significantly affected by drainage quality. Balsam fir exhibited higher foliar toxic secondary compounds concentrations than white spruce in all drainage classes, resulting in lower male pupal mass, survival and longer male developmental time. This, however, did not prevent spruce budworm from consuming more foliage in balsam fir than in white spruce. This response suggests that either natural levels of measured secondary compounds do not provide sufficient toxicity to reduce defoliation, or spruce budworm has developed compensatory mechanisms, which allow it to utilize food resources more efficiently or minimize the toxic effects that are produced by its host's defensive compounds. Larvae exhibited lower pupal mass and higher mortality in rapidly drained and subhygric sites. Drainage class also affected the amount of foliage destroyed but its impact varied over the years and was probably influenced by climatic variables. These results demonstrate the complexity of predicting the effect of resource availability on tree defenses, especially when other confounding environmental factors can affect tree resource allocation and utilization.     

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.     

How does synchrony with host plant affect the performance of an outbreaking insect defoliator?

Phenological mismatch has been proposed as a key mechanism by which climate change can increase the severity of insect outbreaks. Spruce budworm (Choristoneura fumiferana) is a serious defoliator of North American conifers that feeds on buds in the early spring. Black spruce (Picea mariana) has traditionally been considered a poor-quality host plant since its buds open later than those of the preferred host, balsam fir (Abies balsamea). We hypothesize that advancing black spruce budbreak phenology under a warmer climate would improve its phenological synchrony with budworm and hence increase both its suitability as a host plant and resulting defoliation damage. We evaluated the relationship between tree phenology and both budworm performance and tree defoliation by placing seven cohorts of budworm larvae on black spruce and balsam fir branches at different lags with tree budburst. Our results show that on both host plants, spruce budworm survival and pupal mass decrease sharply when budbreak occurs prior to larval emergence. By contrast, emergence before budbreak decreases survival, but does not negatively impact growth or reproductive output. We also document phytochemical changes that occur as needles mature and define a window of opportunity for the budworm. Finally, larvae that emerged in synchrony with budbreak had the greatest defoliating effect on black spruce. Our results suggest that in the event of advanced black spruce phenology due to climate warming, this host species will support better budworm survival and suffer increased defoliation.