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.     

The potential role of ectomycorrhizal fungi in determining Douglas-fir resistance to defoliation by the western spruce budworm (Lepidoptera: Tortricidae)

There is phenotypic variation among individual trees of interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) in their resistance to defoliation by the western spruce budworm (Choristoneura occidentalis Freeman). We evaluated the potential role of ectomycorrhizal fungi in determining this resistance using half-sib seedlings derived from parent trees that are resistant versus susceptible to budworm defoliation in the field. The seedlings were inoculated with Laccaria bicolor ectomycorrhizal fungi, fertilized, or untreated. Approximately 48 d after treatment, late-instar larvae from a nondiapausing laboratory colony of C. occidentalis were allowed to feed on pairs of resistant versus susceptible seedlings for 1 wk. Chemical analyses of current-year shoots for nitrogen (N), phosphorus (P), magnesium (Mg), and zinc (Zn) indicated that the fungus increased foliar concentrations of P and Mg in resistant seedlings, but it did not increase their growth rate. However, L. bicolor had no effect on foliar concentrations of P or Mg in susceptible seedlings, even though seedling growth rates increased slightly in response to the inoculation. L. bicolor had no effect on foliar levels of N or Zn in any of the seedlings. As expected, fertilization increased levels of N and P in the foliage of both resistant and susceptible seedlings, but it did not affect levels of Mg and Zn. Surprisingly, the fertilizer treatment had no effect on seedling growth rates. Despite these differences, late-instar budworms showed no feeding preference among untreated, mycorrhizal, or fertilized seedlings. The fact that seedlings from resistant versus susceptible Douglas-firs responded differently to the L. bicolor treatment lends preliminary support to the hypothesis that ecotmycorrhizae might play a role in Douglas-fir resistance to damage from the western spruce budworm. Finally, differences in foliar concentrations of N and P among untreated seedlings from different maternal trees suggested that foliar nutritional chemistry is influenced by the tree's genotype.     

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.     

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.     

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.     

Larval experience induces adult aversion to rearing host plants: a novel behaviour contrary to Hopkins' host selection principle

1. Hopkins' host selection principle (HSP) states that insects should prefer foliage from their rearing host plant over that of an alternative host. 2. The current study tested whether eastern spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), that were laid and developed on, respectively, resistant and susceptible white spruce Picea glauca (Moench) Voss, showed differences in their feeding and oviposition preferences for these two hosts. 3. The data revealed that previous experience of spruce budworm on a host tree type does not influence the host acceptance and feeding behaviour of later larval stages. However, adult budworm reared on resistant white spruce needles preferentially selected susceptible white spruce needles as the host for their progeny, whereas those reared on susceptible needles showed no preference. 4. To the authors' knowledge, this is the first report of an insect showing an oviposition preference for the non‐rearing host plant. This would tend to increase mixing between insects from susceptible and resistant trees. The present results thus argue against Hopkins' HSP and suggest that learned aversion to resistant foliage experienced by larvae is carried into the adult stage.     

Spruce budworm (Choristoneura spp.) biotype reactions to forest and climate characteristics

The spruce budworm (Choristoneura fumiferana) is the most destructive insect defoliator of forests in North America. Climatic influences on this species' life history are considered a major factor in restricting the extent and intensity of outbreaks. We examine the life history traits of the spruce budworm and related Choristoneura populations with respect to forecasting the conifer‐feeding responses of these insects in changing environments. Analysis of the evolutionary relationships between Choristoneura entities, including their hybridization, genetic distances, and their degree of sympatry leads us to distinguish 15 possible Choristoneura‘biotypes’. Population trend has been associated with recruitment to the feeding stage, and two indicators of recruitment, egg weights and phenological development, are both ‘biotype’ and climate dependent. Among Abietoid feeding ‘biotypes’ and among spruce budworm populations, those from locations with extreme winters tend to have heavier eggs than those from the more benign environments. In spruce budworm, this genetically based adaptation allows populations to increase their potential recruitment substantially where winters are mild. All biotypes feed on the newly developed shoots of their host trees in spring, and are thus vulnerable to the uncertain timing of budbreak. Genetic control of spring emergence is weak so larvae from a single family typically exit from hibernacula over a prolonged period. This guarantees some synchronization with budburst. However, hybrid populations have high heritabilities. This allows rapid adaptation to new conditions (e.g. mixed host‐species stands). Geographic variation in phenological development after establishing feeding sites is largely genetically controlled. The importance of variation in these traits is examined with respect to competing population dynamics theories to evaluate their utility in forecasting future trends in defoliation. We finish with a plea for jointly using alternative approaches in forecasting spatiotemporal patterns of 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.     

Behaviors of Western Spruce Budworm Moths (Choristoneura occidentalis) as Defences Against Bat Predation

We investigated potential defense behaviors of adult western spruce budworm (Choristoneura occidentalis), a non-auditive lepidopteran, against bat predation. Although western spruce budworm moths started to fly before sunset, earlier than many species of moths, temporal isolation of flying moths from foraging bats was incomplete as moths were most active after sunset once bats were foraging. Flying C. occidentalis were most active close to their host trees, and thus were isolated from some bat activity because vegetation limits foraging by some bats. Moths mostly flew near the tops of trees, an area that may have a high predation pressure from bats. Resting western spruce budworm spent little time fluttering their wings or crawling, behaviors that are used as cues by gleaning bats. The outbreak nature of this species, in which large numbers of moths are active at one time, may allow dilution effects to reduce predation risk.     

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.     

Phenological variation as protection against defoliating insects: the case of<Emphasis Type="Italic"> Quercus robur</Emphasis> and<Emphasis Type="Italic"> Operophtera brumata</Emphasis>

Phenological synchrony between budburst and emergence of larvae is critical for the fitness of many spring-feeding insect herbivores. Therefore, large intraspecific variation in timing of budburst of the host may have a negative effect on the herbivore. We studied how asynchrony between emergence of larvae and budburst affects the fitness of Operophtera brumata (Lepidoptera: Geometridae), a major defoliator of Quercus robur, which can adapt to the phenology of a single tree. It is known that, in maturing leaves of Q. robur, accumulation of condensed tannins has a negative effect on growth of O. brumata. However, there is no information about the effect of hydrolysable tannins and other phenolics that are potential antifeedants. In this study, we also analysed changes in secondary chemistry of the foliage of Q. robur and how different compounds are correlated with growth and survival of O. brumata. The effect of asynchrony on O. brumata was studied in rearing experiments. The neonate larvae were incubated without food for different periods of time. The decline in nutritional quality of foliage was estimated by rearing cohorts of larvae with manipulated hatching times on the leaves of ten individual Q. robur trees. For the chemical analysis, the foliage of these trees was sampled at regular intervals. In the absence of foliage, mortality of neonate larvae started to increase exponentially soon after the larvae emerged. If the larvae missed budburst, the decline in nutritional quality of the foliage led to increased mortality and lower body mass (= fecundity). Hydrolysable tannins were not significantly correlated with performance of the larvae. Only condensed tannins were found to correlate negatively with the growth and survival of O. brumata. Certain individual trees were unsuitable hosts for O. brumata because the decline in quality of the foliage was very rapid. Based on regression equations for increasing rate of mortality and decreasing fecundity, we calculated that a relatively small mismatch of +/-30 degree days between budburst and hatching of larvae leads to a 50% decrease in the fitness of O. brumata. Thus, large phenological variation within a Q. robur stand can limit the colonisation of neighbouring trees by dispersing larvae. Furthermore, the hybridisation of moths adapted to phenologically different trees may lead to maladapted phenology of their offspring.     

Determinants of fruit and seed set in Pavonia dasypetala (Malvaceae)

Summary Measurements of foliage quality, physiological, and phenological condition of sample trees were used as independent variables in multiple correlation analyses to determine their effect on female and male spruce budworm larval dry weights. Female budworm from trees high in foliar concentrations of beta-pinene, myrcene and total nitrogen weighed less than those from trees lacking these characteristics. Male budworm from trees high in foliar concentrations of alpha-pinene, myrcene, terpinolene, citronellyl acetate, and bornyl acetate weighted less than those from trees lacking these characteristics. Additionally, relatively vigorous and productive trees tended to be less susceptible (as evidenced by reduced larval weight) to budworm of either sex.     

The role of sugars in western spruce budworm nutritional ecology

Abstract.

• 1 The western spruce budworm, Choristoneura occidentalis Freeman, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, have been used to test the hypothesis that variation in levels of foliar sugars form part of the basis for plant resistance to herbivore attack.
• 2 Budworm population growth was evaluated on artificial diets with 2–45% sucrose using a three generation bioassay. Diets with 1.2% and 3.9% N were tested to determine if responses to sugar were dependent on levels of protein. The 3.9% N diets were supplemented with a mineral salt mixture, so they had high levels of N and minerals.
• 3 The response of budworm population growth to sucrose concentrations ≤20% was convex at 1.2% N and flat for 3.9% N. Population growth on the 1.2% N diet, which had levels of N and minerals similar to host foliage, was good with only 2% sugar, but optimal at the 6% sucrose level; the number of F₁, F₂ and F₃ larvae produced declined substantially when sugar was increased to 11% and 20%. On the 3.9% N diets, population growth was equivalent for diets with 2% and 11% sucrose. Sucrose concentrations ≥29% were detrimental to the budworm at both N levels.
• 4 Sugar concentrations in Douglas-fir foliage varied between 5.7% and 18.4%. Thus, results from the 1.2% N experiment indicated that budworm performance was best on diets with sugar concentrations near the lower limit observed for host foliage. This implies that plants with higher foliar sugar may be inferior hosts for the budworm. Field observations supported this conclusion, as putatively resistant Douglas-fir trees had significantly higher levels of sugars in their foliage than nearby paired susceptible trees. Variation in foliar sugars among individual trees may be part of the mechanism in Douglas-fir resistance to C.occidentalis damage.

     

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.     

The phenological window for western spruce budworm: seasonal decline in resource quality

• 1 Western spruce budworm Choristoneura occidentalis Free. (Lepidoptera: Tortricidae) emerge in the spring before budburst and then face a rapidly deteriorating host quality each season.
• 2 Measures of fitness, survival and fecundity, were made on cohorts of final‐instar spruce budworms deployed on host trees at several times during the season in four field locations in coastal and interior British Columbia, Canada.
• 3 Survival and fecundity were strongly correlated throughout the season and varied as much as four‐fold from maxima at mid‐season to minima at the end of the season.
• 4 Fitness values overall were greatest in the coastal compared with interior locations. Among interior locations, fitness was greatest at the highest elevation and least at the lowest elevation. Both cohort and sample‐based estimates of survival of wild, final‐instar budworms were relatively high in these outbreak populations.
• 5 The influence of the phenological window and degree of synchrony with the host plant on herbivore abundance often depends on other processes affecting population rates of change.

     

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.     

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.     

Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

White pine weevil performances in relation to budburst phenology and traumatic resin duct formation in Norway spruce

1 As the phenological window hypothesis was reported to be significant in influencing the fitness of many herbivores feeding on tree foliage, could it also explain the performance of an insect such as the white pine weevil Pissodes strobi mainly attacking the bark phloem of conifers? 2 Under field conditions, adult weevils were caged on Norway spruce trees presenting a natural variation in their shoot growth phenology. 3 We evaluated white pine weevil biological performances, including oviposition, the number of emerged insects, survival, adult mean weight and tree defense responses as reflected by the production of induced resin canals. 4 None of the white pine weevil biological parameters was significantly affected by Norway spruce phenology. 5 The number of eggs per hole, the number of oviposition holes per leader, the number of emerged adults and their mean weight were not affected by host phenology. 6 The intensity of the traumatic response observed was variable and not correlated with budburst phenology. 7 Trees with higher traumatic responses, forming two or more layers of traumatic ducts, had lower adult emergence and estimated survival. 8 The distance between the first layer of traumatic resin ducts and the start of the annual ring was not correlated with the number of emerged weevils. 9 Norway spruce, which is an exotic tree in North America and a relatively recent host for the white pine weevil, might not possess the defense mechanisms necessary to fight off the white pine weevil.     

Two sides of a coin: host‐plant synchrony fitness trade‐offs in the population dynamics of the western spruce budworm

Conifer‐feeding budworms emerge from overwintering sites as small larvae in early spring, several days before budburst, and mine old needles. These early‐emerging larvae suffer considerable mortality during this foraging period as they disperse in search of available, current‐year buds. Once buds flush, surviving budworms construct feeding shelters and must complete maturation before fresh host foliage senesces and lignifies later in the summer. Late‐developing larvae suffer greater mortality and survivors have lower fecundity when feeding on older foliage. Thus, there is a seasonal trade‐off in fitness associated with host synchrony: early‐emerging budworms have a greater risk of mortality during spring dispersal but gain better access to the most nutritious foliage, while, on the other hand, late‐emerging larvae incur a lower risk during the initial foraging period but must contend with rapidly diminishing resource quality at the end of the feeding period. We investigate the balance that results from these early‐season and late‐season synchrony fitness trade‐offs using the concept of the phenological window. Parameters associated with the variation in the phenological window are used to estimate generational fitness as a function of host‐plant synchrony. Because defoliation modifies these relationships, it is also included in the analysis. We show that fitness trade‐offs characterizing the phenological window result in a robust synchrony relationship between budworm and host plant over a wide geographic range in southern British Columbia, Canada.