Search

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.

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 Quebec. 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.

To determine which ecophysiological factors appear to control monoterpene concentrations in balsam fir foliage [Abies balsamea (L.) P. Mill.], the percentage of photosynthetically active radiation (%PAR), specific leaf area (SLA), light-saturated photosynthesis (A max), and concentrations per unit leaf area of foliar nitrogen (N), total soluble sugars (TSS), starch and monoterpenes were measured on current-year needles from three canopy levels (upper, middle and lower) the year following a pre-commercial thinning. The thinning only modestly changed the light profile within the canopy. %PAR was negatively correlated with SLA (r 2=0.62 in June, r 2=0.53 in July and August) and positively correlated with foliar nitrogen concentrations (r 2=0.51) within the crown profile. The positive relationship between N and A max was quite weak (r 2=0.15), suggesting significant variations in non-photosynthetic N within the canopies. Total monoterpenes were positively correlated with both %PAR (r 2=0.29) and A max (r 2=0.27), and negatively correlated with SLA (r 2=0.30). Contrary to that predicted by the carbon-nutrient balance hypothesis, total monoterpenes were negatively and only very weakly correlated with the starch/N ratio (r 2=0.06) and were not significantly correlated with either the TSS/N or the [TSS+starch]/N ratios. Monoterpenes were positively correlated with both N and TSS, although the relationship varied with the phenological state of the foliage, i.e., monoterpenes were more highly correlated with TSS (r 2=0.67) (immature foliage) in June, and in July and August with N (r 2=0.63) (mature foliage). Thus, it appears that monoterpene concentrations may be controlled primarily by carbohydrate supply in the early growing season and later by enzymatic capacity. Data expressed on a dry weight basis showed a similar pattern.

The leaf area to sapwood area ratio ( Al:As) of trees has been hypothesized to decrease as trees become older and taller. Theory suggests that A l :A s must decrease to maintain leaf-specific hydraulic sufficiency as path length, gravity, and tortuosity constrain whole-plant hy- draulic conductance. We tested the hypothesis that A l :A s declines with tree height. Whole-tree Al:As was measured on 15 individuals of Douglas-fir ( Pseudotsuga menziesii var. menziesii) ranging in height from 13 to 62 m (aged 20-450 years). A l :A s declined substantially as height in- creased (P=0.02). Our test of the hypothesis that A l :A s declines with tree height was extended using a combina- tion of original and published data on nine species across a range of maximum heights and climates. Meta-analysis of 13 whole-tree studies revealed a consistent and signif- icant reduction in A l :A s with increasing height (P

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing indicated that the feeding behaviour of the larvae, which is affected by the insect population density, significantly influenced the impact of balsam fir, Abies balsamea, staminate flowering on spruce budworm biology. At low budworm density, the production of pollen in the midcrown of host trees reduced the insect development time by 5 days without affecting pupal weight, fecundity and survival. However, at high budworm density, the small amount of current-year foliage produced by flowering branches forced old larvae (sixth instar) either to feed on 1-year-old foliage (backfeeding) or to move from the midcrown to the lower crown section where staminate flowers are absent and more current-year foliage is available. When old larvae fed on old foliage, they exhibited reduced pupal weight and fecundity without losing the advange in development time that they obtained from feeding on pollen during their early stages of development. On the other hand, when old larvae moved to the lower crown section, they avoided the negative effects of backfeeding but lost the advantage in development time that was gained from feeding on pollen. Results from this study indicated that the production of staminate flowers by balsam fir trees could have opposite effects on spruce budworm population dynamics depending upon the insect population density when flowering occurs.

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.

Two field studies were carried out to determine the influence of Abies balsamea foliage age on the preference and performance of larvae of Neodiprion abietis, a specialist Diprionid sawfly. Preference was determined by examining N. abietis defoliation on all age classes of foliage. Performance was estimated using larval survival, cocoon weights and the percentage of adults that were females. Neodiprion abietis preference for, and performance on, current-year foliage was very low, peaked on 2 or 3-year-old foliage, and declined on older foliage. Thus, sawfly feeding preference was adaptive. However, survival and cocoon weight were highest when sawflies were allowed to feed on all age classes of foliage, demonstrating that an insect specialist may perform better when feeding on several age classes of foliage from a single host plant species. These results indicate that either different larval instars have different nutritional requirements, or that food mixing provides the best diet, permitting the herbivore to obtain needed nutrients while avoiding ingestion of toxic doses of secondary metabolites. In addition, our results suggest that limited availability of varied foliage has more negative consequences for N. abietis females than for males, as the percentage of survivors that were females decreased when juvenile mortality was high. Our results emphasize the importance of considering non-linear changes in foliar quality as leaves age on herbivore preference and performance, and demonstrate how a herbivore can use this variability to maximize its fitness.

Stand thinning across different soil drainage types was used to test the ecophysiological basis for the control of foliar monoterpenes in current-year foliage of balsam trees [Abies balsamea (L.) P. Mill.] the year following treatment. Photosynthetic capacity (Amax) was greater on mesic sites than on sub-hygric and hydric sites (1.81, 1.33 and 0.88 µmol m–2 s–1, respectively) and greater in mid-July than at the end of June (1.86 vs 0.77 µmol m–2 s–1). Foliar N was greater on thinned plots than control plots (2.6 vs 2.2 g m–2); starch was greater on mesic control plots than mesic thinned plots (16.3 vs 10.9 g m–2); and total soluble sugars (TSS) were higher in mid-July than at the end of June (16.5 vs 12.8 g m–2). Total monoterpenes, as well as five individual monoterpenes, showed a complex interaction between site drainage class, thinning treatment and time of sampling (P

Moose were observed to browse preferentially on balsam fir (Abies balsamea) trees in stands where the stem density had been mechanically reduced to about 2000 stems/ha from over 30,000 stems/ha. Twigs from trees in thinned and unthinned stands were analyzed to test the hypotheses that moose were choosing thinned stands to maximize intake of a nutrient, or avoiding plant secondary compounds deliterious to digestion. Analyses included: twig length, weight, diameter, acid detergent fiber, neutral detergent fiber, dry matter digestibility, macroelements, crude fat (resins), crude protein, ash, and volatile secondary metabolites. Significantly higher concentrations of crude protein, P, Ca, Na, and crude fats occurred in trees from thinned compared to unthinned stands. Twigs from thinned stands were more digestible, longer, heavier, and had a greater diameter than those from unthinned stands. Several secondary metabolites were found in highest concentration in thinned stands. We suggest that moose chose thinned stands over unthinned stands for feeding because of high protein levels and large twigs. Trees in unthinned stands had protein levels below reported maintenance levels required by moose. Secondary metabolite concentrations were opposite to the direction predicted and the data do not support plant defence hypotheses for the chemicals analyzed.

The hypothesis that annual climatic variation can affect the nutritive value of host-plant tissue for a herbivore was tested. Larvae of a single generation of western spruce budworm, Choristoneura occidentalis Free., were fed foliages that had been collected on five consecutive springs from a stand of balsam fir Abies balsamea (L.) Mill. Budworm of both sexes were about 20 percent heavier at pupation when fed folages that had been produced in early rather than later spring seasons. This could result in an estimated 25 percent difference in the reproductive rate of females. It is concluded that annual variations in spring climate can affect the nutritive value of forage.