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.     

Cross‐scale effects of spruce budworm outbreaks on boreal warblers in eastern Canada

Insect outbreaks are major natural disturbance events that affect communities of forest birds, either directly by affecting the food supply or indirectly by changing the vegetation composition of forest canopies. An examination of correlations between measures of bird and insect abundance across different spatial scales and over varying time lag effects may provide insight into underlying mechanisms. We developed a hierarchical Bayesian model to assess correlations between counts of eight warbler species from the Breeding Bird Survey in eastern Canada, 1966 to 2009, with the presence of spruce budworm (Choristoneura fumiferana Clem.) at immediate local scales and time‐lagged regional scales, as measured by extent of defoliation on host tree species. Budworm‐associated species Cape May warbler (Setophaga tigrina), bay‐breasted warbler (Setophaga castanea), and Tennessee warbler (Oreothlypis peregrina) responded strongly and positively to both local and regional effects. In contrast, non‐budworm‐associated species, Blackburnian warbler (Setophaga fusca), magnolia warbler (Setophaga magnolia), Canada warbler (Cardellina canadensis), black‐throated blue warbler (Setophaga caerulescens), and black‐throated green warbler (Setophaga virens), only responded to regional effects in a manner that varied across eastern Canada. The complex responses by forest birds to insect outbreaks involve both increased numerical responses to food supply and to longer term responses to changes in forest structure and composition. These effects can vary across spatial scales and be captured in hierarchical population models, which can serve to disentangle common trends from data when examining drivers of population dynamics like forest management or climate change.     

Effects of artificial and western spruce budworm (Lepidoptera: Tortricidae) defoliation on growth and biomass allocation of Douglas-fir seedlings

Artificial defoliation has been used commonly to simulate defoliation by insect herbivores in experiments, in spite of the fact that obvious differences exist between clipping foliage and natural defoliation due to insect feeding. We used a greenhouse experiment to compare the effects of artificial and western spruce budworm (Choristoneura occidentalis Freeman) defoliation on the growth and biomass allocation of 3-yr old half-sib seedlings from mature Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco variety glauca] trees that showed phenotypic resistance versus susceptibility to budworm defoliation in the forest. Artificial clipping of buds mimicked the effects of budworm feeding on total seedling biomass when 50% of the terminal buds were damaged. However, artificial defoliation decreased seedling height, relative growth rate of height, and shoot: root ratio more than budworm defoliation, whereas budworm defoliation decreased stem diameter relative growth rate more than artificial defoliation. Half-sib seedling progeny from resistant maternal tree phenotypes had greater height, diameter, biomass, and shoot: root ratio than seedlings from susceptible phenotypes. We concluded that careful artificial defoliation could generally simulate effects of budworm defoliation on total biomass of Douglas-fir seedlings, but that the two defoliation types did not have equal effects on biomass allocation between shoot and root. Further, an inherently higher growth rate and a greater allocation of biomass to shoot versus root are associated with resistance of Douglas-fir trees to western spruce budworm defoliation.     

Impacts of spruce budworm defoliation on the habitat of woodland caribou,moose, and their main predators

Forest logging has contributed to the decline of several woodland caribou populations by causing the fragmentation of mature coniferous stands. Such habitat alterations could be worsened by spruce budworm (SBW) outbreaks. Using 6201 vegetation plots from provincial inventories conducted after the last SBW outbreak (1968–1992) in boreal forests of Québec (Canada), we investigated the influence of SBW‐caused tree defoliation and mortality on understory vegetation layers relevant to woodland caribou and its main predators. We found a positive association between severe outbreaks and the cover of most groups of understory plant species, especially in stands that were dominated by balsam fir before the outbreak, where a high canopy openness particularly benefited relatively fast‐growing deciduous plants. Such increases in early successional vegetation could provide high‐quality forage for moose, which is likely to promote higher wolf densities and increase predation pressure on caribou. SBW outbreaks may thus negatively affect woodland caribou by increasing predation risk, the main factor limiting caribou populations in managed forests. For the near future, we recommend updating the criteria used to define critical caribou habitat to consider the potential impacts of spruce budworm defoliation.     

Interguild interactions between a non-native phloem feeder and a native outbreaking defoliator

1. Competitive and synergistic interactions directly or indirectly drive community dynamics of herbivorous insects. Novel interactions between non-native and native insects are unpredictable and not fully understood. 2. We used manipulative experiments on mature red spruce trees to test interactions between a non-native phloem feeding insect, the brown spruce longhorn beetle (BSLB), and an outbreaking native defoliator, the spruce budworm. We subjected treatment trees to defoliation by three densities of spruce budworm larvae. Treatment trees were: stressed by (i) girdling (to mimic beetle feeding) or (ii) girdling + BSLB before spruce budworm larvae were introduced on branches in sleeve cages. Budworm larvae then fed on foliage and developed to pupation. We assessed all branches for budworm performance, defoliation, shoot production and shoot growth. 3. Shoot length did not differ in response to stress from girdling or BSLB infestation. Neither stress from girdling, nor interactions with BSLB feeding affected spruce budworm performance or defoliation. Intraspecific impacts on performance and defoliation in relation to budworm density were stronger than the effects of tree stress. 4. Prior infestation of red spruce by BSLB in our experimental set-up did not influence spruce budworm performance. BSLB is a successful invader that has blended into its novel ecological niche because of ecological and phylogenetic similarities with a native congener, Tetropium cinnamopterum. 5. Outbreaks by BSLB will not likely impede or facilitate spruce budworm outbreaks if they co-occur. It would be useful to evaluate the reverse scenario of BSLB success after defoliation stress by spruce budworm.     

Ecological mechanisms of population change during outbreaks of the spruce budworm

Abstract.  1. Stage-specific survival and recruitment of spruce budworm were measured by frequent sampling of foliage in four outbreak populations over a 15-year period in Ontario and Quebec, Canada.
2. Patterns of change in population density during the outbreak collapse phase were closely linked to changes in survival of the late immature stages, and were determined largely by the impact of natural enemies.
3. Host-plant feedback also contributed significantly to survival patterns throughout the outbreak: annual defoliation influenced survival of fourth and fifth instars and fecundity while cumulative defoliation influenced survival of the very early larval stages (first and second) via impacts on stand condition.
4. Inclusion of this host-plant feedback reveals spruce budworm population dynamics as a function of density-related trophic interactions that vary in their order and strength of influence over time. This view re-introduces the importance of forest interactions as a component of dynamics of the spruce budworm.     

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.     

A lagged, density-dependent relationship between jack pine budworm Choristoneura pinus pinus and its host tree Pinus banksiana

Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.     

Incidence of mesoscale convergence lines as input to spruce budworm control strategies

On occasion Greenbank et al. (1980) in their 1973 to 1976 study on spruce budworm moth dispersal in New Brunswick, Canada, detected intense line concentrations of airborne moths crossing special radar observing sites located about 100 km inland from both the Bay of Fundy and the Northumberland Straits. Line concentrations of insects reflect the presence of atmospheric convergence lines. Data from a surface mesonetwork and wind and temperature soundings up to 2 km collected in New Brunswick from 1976 to 1978 around the period of spruce budworm moth activity has revealed sea breeze fronts to be the meteorological origin for the line concentrations of moths. Analysis has shown sea breeze fronts penetrating 80 to 100 km inland in New Brunswick can be expected once or twice each year during the moth dispersal period. An equation for maximum inland penetration was developed which gave a correlation coefficient of 0.78 between predicted and observed inland penetrations with a standard error of 17.5 km. Predicted sea-breeze frontal penetrations of 100 km or more were considered likely to produce a line concentration of moths observable at the radar sited. Using this criteria the prediction scheme delineated four out of the five nights when line concentrations of moths were observed and only over-predicted on one occasion. Thus meteorological data can be used to predict the appearance of line concentrations of spruce budworm moths and so provide input into population redistribution studies and into the development of control strategies directed at the adult budworm.Presented at Eighth International Congress in Biometeorology, 9–14 September 1979, Shefayim, Israel.     

Variation in budburst phenology of Douglas-fir related to western spruce budworm (Lepidoptera: Tortricidae) fitness

Variation in budburst phenology among individual trees of interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) may influence their susceptibility to western spruce budworm (Choristoneura occidentalis Freeman) defoliation. We tested the hypothesis that phenological asynchrony between Douglas-fir and the western spruce budworm is a mechanism of resistance using clones derived from parent trees that showed resistance versus susceptibility to C. occidentalis defoliation in the field. Susceptible clones had earlier budburst phenology compared with resistant clones when they were grown in a common greenhouse environment, demonstrating a genetic basis for parallel phenological differences exhibited by the parent trees. We tested the importance of phenological asynchrony as a factor influencing fitness of C. occidentalis using two different greenhouse bioassay experiments. One experiment compared western spruce budworm performance on equivalent phenological stages of susceptible and resistant clones by matching larval feeding to the columnar (fourth) bud development stage of each clone. Larvae reared on resistant clones had greater realized fitness (i.e., number of F1 offspring produced) than those reared on susceptible clones when the influence of variation in budburst phenology was minimized. In the other experiment, western spruce budworm larvae were placed on all trees on the same date when approximately 50% of all terminal buds in the population were in the yellow (second) budburst stage. Larvae reared on susceptible clones had greater realized fitness than those reared on resistant clones when the influence of phenological asynchrony was expressed. Our results suggest that resistant phenotypes of Douglas-fir have negative effects on survival and reproduction of C. occidentalis under the natural conditions that insects and trees experience in the field. Genetic variation among trees in budburst phenology has an important influence on interactions between the western spruce budworm and Douglas-fir.     

Dendroentomological and forest management implications in the Interior Douglas-fir zone of British Columbia, Canada

Douglas-fir (Pseudotsuga menziesii var. glauca Mirb. Franco) forests in the Interior of British Columbia, Canada, show periodic defoliation due to western spruce budworm (WSB) (Choristoneura occidentalis Freeman) outbreaks. Tree defoliation causes a reduction in radial growth and is therefore visible in tree rings. In this paper, we identify WSB defoliation history, and critically examine the potential for using dendrochronological analysis by comparing tree-ring estimates with insect surveys. WSB defoliation history was investigated using cores from Douglas-fir growing in the Lac du Bois region of the Kamloops Forest District. Years with an abrupt decrease in radial growth were considered as negative pointer years that potentially reflected WSB outbreaks. The comparison with ponderosa pine (Pinus ponderosa Dougl., ex P. & C. Laws.) (a non-host species) permitted differentiation between growth reductions in Douglas-fir due to climatic effects and those due to defoliation by WSB. The dendrochronological data were matched with information reporting visible damage in Forest Insect Disease Survey (FIDS) and British Columbia Ministry of Forest records. Our objective-based method using ring-width measurements from host and non-host chronologies was compared with qualitative techniques based on the software program OUTBREAK. We were able to distinguish seven distinct outbreak events in 300 years of record.     

Identifying spatial relationships at multiple scales: principal coordinates of neighbour matrices (PCNM) and geostatistical approaches

We compared two methodological approaches – principal coordinate analysis of neighbour matrices (PCNM) and geostatistics – that both aim at extracting several spatial scales in order to identify spatial relationships between organisms and environmental variables at multiple scales. From a statistical point of view, PCNM analysis and geostatistics come from "two different worlds"– PCNM is based on classical "data analysis" while geostatistical modelling is developed in a probabilistic context. These two methods were used to investigate the spatial relationships between defoliation caused by spruce budworm Choristoneura fumiferana and bioclimatic conditions in Ontario since 1941 through a wide range of scales. On the one hand, PCNM variables related to defoliation frequency were partitioned into four spatial submodels representing respectively four spatial scales: very broad scale (ca>300 km), broad scale (ca 180 km), fine (ca 100 km), and very fine (<80 km). On the other hand, nested variogram modelling was used to identify the relevant scales. The nested variogram model was composed of four variograms with different characteristic scales close to those of the PCNM spatial submodels. Maps of PCNM submodels and kriging components revealed similar spatial patterns of defoliation frequency at very broad and broad scales while spatial patterns at fine and very fine scales looked quite different. Both methods showed that defoliation by spruce budworm occurs at the broader spatial scales but may be explained by fluctuations at the smaller scales. Finally, results based on geostatistics using a Linear Model of Coregionalisation suggested that climatic conditions can be considered to act at the level of outbreak dynamics while the tree community of spruce budworm's principal hosts controls local population dynamics.     

Taste receptor activity and feeding behaviour reveal mechanisms of white spruce natural resistance to Eastern spruce budworm Choristoneura fumiferana

The pattern of feeding of Eastern spruce budworm Choristoneura fumiferana (Clem.) (Lepidoptera, Tortricidae) is compared on foliage from white spruce Picea glauca (Moench) Voss. (Pinaceae) trees previously determined to be susceptible and resistant to defoliation by budworm. No differences are observed in electrophysiological responses from taste sensilla to aqueous extracts of the two foliage types, nor is there a preference for either extract type in a choice test. Acetone extracts from the two foliage types are both preferred to a control sucrose solution, although neither elicits a preference relative to the other. These results suggest that there is no difference in phagostimulatory power of internal leaf contents of the two foliage types. Longer‐term observation of feeding behaviour in a no‐choice situation shows no difference in meal duration, confirming the lack of difference in phagostimulatory power. However, on average, intermeal intervals are twice as long on the resistant foliage, leading to an overall lower food consumption during the assay. This result suggests an anti‐digestive or toxic effect of the resistant foliage that slows behaviour and limits food intake. Previous research has shown that waxes of the resistant foliage deter initiation of feeding by the spruce budworm and that this foliage contains higher levels of tannins and monoterpenes. The data suggest that the resistant foliage contains a post‐ingestive second line of defence against the spruce budworm.     

Dendroentomological and forest management implications in the Interior Douglas-fir zone of British Columbia,Canada

Douglas-fir (Pseudotsuga menziesii var. glauca Mirb. Franco) forests in the Interior of British Columbia, Canada, show periodic defoliation due to western spruce budworm (WSB) (Choristoneura occidentalis Freeman) outbreaks. Tree defoliation causes a reduction in radial growth and is therefore visible in tree rings. In this paper, we identify WSB defoliation history, and critically examine the potential for using dendrochronological analysis by comparing tree-ring estimates with insect surveys. WSB defoliation history was investigated using cores from Douglas-fir growing in the Lac du Bois region of the Kamloops Forest District. Years with an abrupt decrease in radial growth were considered as negative pointer years that potentially reflected WSB outbreaks. The comparison with ponderosa pine (Pinus ponderosa Dougl., ex P. & C. Laws.) (a non-host species) permitted differentiation between growth reductions in Douglas-fir due to climatic effects and those due to defoliation by WSB. The dendrochronological data were matched with information reporting visible damage in Forest Insect Disease Survey (FIDS) and British Columbia Ministry of Forest records. Our objective-based method using ring-width measurements from host and non-host chronologies was compared with qualitative techniques based on the software program OUTBREAK. We were able to distinguish seven distinct outbreak events in 300 years of record.     

Phenological synchrony between eastern spruce budworm and its host trees increases with warmer temperatures in the boreal forest

Climate change is predicted to alter relationships between trophic levels by changing the phenology of interacting species. We tested whether synchrony between two critical phenological events, budburst of host species and larval emergence from diapause of eastern spruce budworm, increased at warmer temperatures in the boreal forest in northeastern Canada. Budburst was up to 4.6 ± 0.7 days earlier in balsam fir and up to 2.8 ± 0.8 days earlier in black spruce per degree increase in temperature, in naturally occurring microclimates. Larval emergence from diapause did not exhibit a similar response. Instead, larvae emerged once average ambient temperatures reached 10°C, regardless of differences in microclimate. Phenological synchrony increased with warmer microclimates, tightening the relationship between spruce budworm and its host species. Synchrony increased by up to 4.5 ± 0.7 days for balsam fir and up to 2.8 ± 0.8 days for black spruce per degree increase in temperature. Under a warmer climate, defoliation could potentially begin earlier in the season, in which case, damage on the primary host, balsam fir may increase. Black spruce, which escapes severe herbivory because of a 2‐week delay in budburst, would become more suitable as a resource for the spruce budworm. The northern boreal forest could become more vulnerable to outbreaks in the future.     

Warming counteracts defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony

Climate change is altering phenology; however, the magnitude of this change varies among taxa. Compared with phenological mismatch between plants and herbivores, synchronization due to climate has been less explored, despite its potential implications for trophic interactions. The earlier budburst induced by defoliation is a phenological strategy for plants against herbivores. Here, we tested whether warming can counteract defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony. We compared the larval phenology of spruce budworm and budburst in balsam fir, black spruce, and white spruce saplings subjected to defoliation in a controlled environment at temperatures of 12, 17, and 22°C. Budburst in defoliated saplings occurred 6–24 days earlier than in the controls, thus mismatching needle development from larval feeding. This mismatch decreased to only 3–7 days, however, when temperatures warmed by 5 and 10°C, leading to a resynchronization of the host with spruce budworm larvae. The increasing synchrony under warming counteracts the defoliation‐induced mismatch, disrupting trophic interactions and energy flow between forest ecosystem and insect populations. Our results suggest that the predicted warming may improve food quality and provide better growth conditions for larval development, thus promoting longer or more intense insect outbreaks in the future.     

Uneven Rates of Landscape Change as a Source of Bias in Roadside Wildlife Surveys

ABSTRACT Roadside survey data have been used frequently to assess species occurrence and population trends and to establish conservation priorities. However, most studies using such data assume that samples are representative of either the amount of habitat or its rate of change at larger spatial scales. We tested both of these assumptions for the Breeding Bird Survey (BBS) from 1974 to 2001 in New Brunswick, Canada. Our study focused on mature forest—a cover type that we predicted would be characterized by rapid change due to human activities and that is of high ecological importance. We also sought to determine whether land cover changes adjacent to BBS routes were related to bird population trends detected in BBS data. Within all 3 time periods examined (1970s, 1980s, and 1990s), the amount of mature forest adjacent to BBS routes was significantly lower than in surrounding 1° blocks of latitude and longitude. This could be problematic for studies that use roadside data to compare the relative abundance of species. On average, mature forest declined at a rate of-1.5% per year over the 28-year study period. We detected no significant difference in the rate of change between degree blocks and BBS routes over this time span. However, in the 1970s and 1980s, mature forest declined more rapidly in degree blocks (-2.7%/yr) than adjacent to BBS routes (-0.5/yr). We also found that the BBS trend for a mature forest-associated species, blackburnian warbler (Dendroica fusca), was correlated with the trend in mature forest along BBS routes. This, combined with slower rates of mature forest change along routes in the 1970s and 1980s, suggests that BBS data may have underestimated population declines during this period. It is important that research be conducted to test for potential biases in roadside surveys caused by uneven rates of landscape change, particularly in regions characterized by rapid habitat alteration.     

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.     

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.