Spruce budworm feeding and oviposition are stimulated by monoterpenes in white spruce epicuticular waxes

Monoterpenes, source of the distinctive odor of conifers, are generally considered plant defensive compounds. However, they are also known to act as long‐range insect attractants, as they are volatile and permeate forest airspaces. Moreover, they are lipid soluble and can be absorbed into plant epicuticular waxes. We test their role in short‐range host plant choice by both adult females and larvae of a folivorous forest pest (Choristoneura fumiferana). We conducted laboratory assays testing the responses of Eastern spruce budworm to an artificial monoterpene mix (α‐pinene, β‐pinene, limonene, myrcene) and to white spruce (Picea glauca) epicuticular waxes in closed arenas. Ovipositing females preferred filter paper discs treated with P. glauca waxes to controls, and preferred the waxes + monoterpenes treatment to waxes alone. However, females showed no preference between the monoterpene‐treated disc and the control when presented without waxes. Feeding larvae prefered wax discs to control discs. They also consumed discs treated with realistic monoterpene concentrations and wax preferentially over wax‐only discs, but showed no preference between extremely high monoterpene concentrations and wax‐only controls. In an insect‐free assay, P. glauca epicuticular wax decreased monoterpene volatilization. These results suggest that P. glauca waxes and realistic concentrations of monoterpenes are stimulatory to both egg‐laying females and feeding larvae, and that their effects are synergistic.     

Larval feeding behaviour affects the impact of staminate flower production on the suitability of balsam fir trees for spruce budworm

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.     

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).     

Oviposition behavior in spruce budwormChoristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

Epicuticular waxes of balsam fir, Abies balsamea,act as oviposition stimulants for fertilized females of the eastern spruce budworm, Choristoneura fumiferana(Clem.). Those same waxes do not stimulate unfertilized females, nor do the latter respond to nonchemical cues such as leaf texture, shape, etc. Ablation experiments suggest that the chemoreceptors responsible for the perception of waxes are located on the proboscis.     

Nutritional physiology of the eastern spruce budworm,Choristoneura fumiferana,infected with Nosema fumiferanae,and interactions with dietary nitrogen

Summary Female eastern spruce budworm larvae, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae), inoculated with a medium lethal spore dosage of the microsporidium Nosema fumiferanae (Thomson) exhibited significant reductions in consumptive index (CI), nitrogen consumptive index (NCI), relative growth rate (RGR), and gross (ECI) and net (ECD) production effeciencies when compared to microsporidian-free larvae. Diseased larvae also exhibited significant increases in approximate digestibility (AD), N utilization efficiency (NUE), and larval moisture content. Both healthy and diseased insects were reared on 2.5% N and 4.5% N diets. Those on the 2.5% N diet showed significant increases in CI, although NCI was still lower than NCI measured for larvae reared on 4.5% N. NUE was also higher on the 2.5% N diet. Diseased cohorts reared on 2.5% N diet had significantly greater mortality than those reared on 4.5% N diet. Pupal weight and development time of infected individuals did not respond to dietary N concentration. However, healthy insects achieved greater pupal weights in a shorter time on the 4.5% N diet than those on the 2.5% N diet. Mortality of healthy insects was unaffected by dietary N.     

Impact of balsam fir flowering on pollen and foliage biochemistry in relation to spruce budworm growth, development and food utilization

The impact of balsam fir (Abies balsamea (L.) Miller) flowering on nutritional and allelochemical quality of pollen, current-year and one-year-old foliage is studied in relation to spruce budworm (Choristoneura fumiferana Clem.) (Lepidoptera: Tortricidae) growth, development and utilization of food and nitrogen. In the laboratory, using fresh food from the field, we simulated conditions of low larval population density, in which there is no current-year foliage depletion during the spruce budworm feeding period. Similarly, we simulated conditions of high larval population density when current-year foliage depletion occurs.Because of the high nutritive value of pollen (high amounts of amino acids and minerals, especially nitrogen; low monoterpene content), insects from flowering trees reached the fifth instar five days earlier than those from non-flowering trees, and had heavier dry- and nitrogen-weights at the beginning of the fifth instar. At budbreak, switching from pollen to current-year foliage negatively affected conversion efficiencies and digestibilities of food and nitrogen (AD; ADN; ECDN; ECI; ECIN). The switch from pollen to new foliage had a detrimental impact on fifth-instar survival and on newly-moulted sixth-instar dry- and nitrogen-weights. Moreover, during the fifth instar, balsam fir flowering reduced the nutritive value of current-year foliage, which in turn, might have contributed to the reduced larval growth. Nevertheless, during the sixth instar, balsam fir flowering affected the biochemistry of current-year foliage in ways that enabled larvae to compensate for their low fifth-instar biological performance; larvae also managed to reach pupal dry weight similar to larvae reared on non-flowering trees. Current-year foliage from flowering trees contained less nitrogen, total soluble sugars and total monoterpenes. Those foliar characteristics enabled larvae to increase food and nitrogen consumption rates (RCR; RNCR), because of lower repellency and/or post-ingestional feedback from monoterpenes.As for current-year foliage, balsam fir flowering reduced nitrogen, total soluble sugar and total monoterpene contents in one-year-old foliage during the sixth-instar feeding period. These characteristics enabled sixth-instar larvae, fed on old foliage from flowering trees, to have high relative food and nitrogen consumption rates (RCR; RNCR). Larvae were also able to reach higher relative growth rates (RGR) and relative nitrogen accumulation rates (RNAR) compared to larvae reared on one-year-old foliage from non-flowering trees. Finally, larvae on flowering trees had pupal dry weight similar to those from non-flowering trees, but reached the adult stage nine days earlier.Regardless the foliage type consumed by spruce budworm larvae during the sixth instar, pollen consumption during early larval stages reduced total development time, and thus exposure time to natural enemies. This phenomenon might increase larval survival. Balsam fir flowering changed the biochemistry of one-year-old and current-year foliages, but did not affect pupal dry weights of larvae reared on flowering trees compared to those reared on non-flowering trees. Thus, at low population density, spruce budworm populations in balsam fir flowering stands might be favoured over those in balsam fir non-flowering stands. In addition, when larvae consumed one-year-old foliage during the entire sixth instar, those on flowering trees are probably favoured over those on non-flowering trees. However, because flowering trees produce less new foliage than non-flowering trees, current-year foliage depletion may occur earlier on flowering trees than on non-flowering trees. Thus, at similar larval population density, larvae on flowering trees might have to feed on one-year-old foliage earlier than those on non-flowering trees. In that case, spruce budworm populations on non-flowering stands would be favoured over those on flowering stands.     

Oral exudate as a mediator of behavior in larval eastern and western spruce budworms (Lepidoptera: Tortricidae)

The production of oral exudate by larval eastern and western spruce budworms,Choristoneura fumiferana (Clem.) andChoristoneura occidentalis Free., respectively, was investigated in the laboratory. All larvae except those entering into a molt exhibited aggressive behavior and produced exudate in response to handling or intraspecific encounters. Larvae could be induced to produce exudate up to four times over 2–3 min and produced an average of 1.92±0.04 µl (X ± SE) per induction. Larvae on foliage spent much of their time maintaining their silken feeding tunnel, including spinning and combing silk and removing frass. Exposure to conspecific oral exudate deposited inside the tunnel, or released by agitated larvae inside the tunnel, increased the proportion of larvae that dispersed away from the tunnels and, apparently, increased the larval sensitivity to disturbances. The behavior induced by the oral exudate indicates that it acts as an epideictic (spacing) pheromone.     

Age-related responses from the maxillary sensilla styloconica of Choristoneura fumiferana larvae to foliage extracts from balsam fir hosts

An electrophysiological study of the sensilla styloconica of the galea in Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae) larvae showed a differential response between fourth- and sixth-instars to extracts of balsam fir foliage. Larvae raised on artificial diet were stimulated with the water soluble fraction of needle extracts obtained from terminal and lateral shoots of 30- and 70-yr-old balsam fir trees. An extract-sensitive neuron was found in the lateral styloconic sensillum of both instars. The lateral styloconica in the fourth-instar larvae were more sensitive to extracts from terminal than from lateral shoot foliage of both young and old trees. The lateral styloconica of sixth-instar larvae were more sensitive to lateral shoot foliage of old trees. Results are discussed with respect to their relationship to feeding preferences and feeding rates observed in a previous behavioural study.     

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.     

Feeding preferences of eastern spruce budworm larvae in two-choice tests with combinations of host-plant extracts

Three extracts were obtained from each of four host plants: sugars/glycosides; amino acids/bases; and organic acids. The four hosts studied were balsam fir, black spruce, red spruce, and white spuce. The effects of recombined host plant extracts on the feeding behaviour of eastern spruce budworm larvae were examined in two-choice tests. Results compare feeding preferences as well as feeding rates between combinations of extracts. The importance of each class of polar compounds is discussed.

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Préférences des larves de la tordeuse des bourgeons de l'épinette envers extraits combinés de leurs plantes-hôtes

Résumé Trois extraits ont été obtenus de chacune de quatre plantes-hôtes: sucres/glycosides; acides aminées/bases organiques; acides organiques. Les quatre plantes-hôtes étudiées furent le sapin baumier, et les épinettes blanc, noir, et rouge. Le comportement gustatoire des larves a été étudié en raison de l'effet de combinaisons d'extraits de chaque plantehôte. Les résultats comparent la préférence des larves ainsi que leur consommation totale envers ces extraits.

     

Persistence of balsam fir and black spruce populations in the mixedwood and coniferous bioclimatic domain of eastern North America

The boreal ecocline (ca 49°N) between the southern mixedwood (dominated by balsam fir) and the northern coniferous bioclimatic domain (dominated by black spruce) may be explained by a northward decrease of balsam fir regeneration, explaining the gradual shift to black spruce dominance. 7,010 sample plots, with absence of major disturbances, were provided by the Quebec Ministry of Forest, Fauna, and Parks. The regeneration (sapling abundance) of balsam fir and black spruce were compared within and between the two bioclimatic domains, accounting for parental trees, main soil type (clay and till) and climate conditions, reflected by summer growing degree‐days above 5°C (GDD_5), total summer precipitation (May–August; PP_MA). Parental trees and soil type determined balsam fir and black spruce regeneration. Balsam fir and black spruce, respectively, showed higher regeneration in the mixedwood and the coniferous bioclimatic domains. Overall, higher regeneration was obtained on till for balsam fir, and on clay soils for black spruce. GDD_5 and PP_MA were beneficial for balsam fir regeneration on clay and till soils, respectively, while they were detrimental for black spruce regeneration. At a population level, balsam fir required at least 28% of parental tree basal area in the mixedwood, and 38% in the coniferous bioclimatic domains to maintain a regeneration at least equal to the mean regeneration of the whole study area. However, black spruce required 82% and 79% of parental trees basal area in the mixedwood and the coniferous domains, respectively. The northern limit of the mixedwood bioclimatic domain was attributed to a gradual decrease toward the north of balsam fir regeneration most likely due to cooler temperatures, shorter growing seasons, and decrease of the parental trees further north of this northern limit. However, balsam fir still persists above this northern limit, owing to a patchy occurrence of small parental trees populations, and good establishment substrates.     

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.     

Food consumption and utilization by 6th-instar larvae of spruce budworm, Choristoneura fumiferana: a comparison on three Picea (spruce) species

The effects of tree species, shoot age, and sex of larvae on food consumption and utilization were investigated in a factorial experiment. Diets prepared from lyophilized and powdered current-year shoots, from two age classes (sample dates) of Picea glauca (Moench) Voss, Picea rubra, Sarg., and Picea mariana (Mill.) B.S.P., were evaluated for eight performance criteria. All factors had significant effects on all criteria except for tree species on assimilation efficiency, shoot age on development time, and sex on net growth efficiency. Eight of the possible 24 two-factor interactions and one of the possible eight three-factor interactions were significant. Food utilization efficiencies and relative growth rate were in the low and moderate characterization limits for immature arthropods; relative consumption rates were high. The higher performance values on Picea glauca were related to this species greater vulnerability to spruce budworm. Performance criteria of similar larvae on an artificial diet were compared with those on the foliage-diets.

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Résumé L'auteur a déterminé la quantité d'aliments utilisées, à 22±0,5 °C, par des larves de Tordeuse des bourgeons de l'épinette parvenues à leur sixième stade de développement qui s'étaient nourries d'un régime artificiel et d'aiguilles de l'année en cours lyophilisées et réduites en poudre, prélevées à deux dates différentes, à 10 jours d'intervalle, et provenant d'épinettes blanchés (Picea glauca [Moench] Voss), d'épinettes rouges (Picea rubra Sarg.) et d'épinettes noires (Picea mariana [Mill.] B.S.P.). Les femelles ont invariablement ingéré une quantité significativement plus importante de tous les régimes que les mâles, consommant 24% de plus d'aiguilles d'épinette noire et 73% de plus d'aiguilles d'épinette blanche.Les femelles prenaient en général plus de temps pour parvenir au stade de chrysalide et avaient un taux d'utilisation des aliments significativement plus élevé. L'efficacité nette de la croissance chez les deux sexes ne présentait toutefois aucune différence significative. La performance des larves ayant ingéré des aiguilles provenant de deux classes d'âge différait de façon significative mais non pas uniforme d'un régime à l'autre: les aiguilles d'épinette blanche plus récentes donnaient une performance généralement plus élevée contrairement aux aiguilles plus jeunes d'épinette rouge et d'épinette noire. Les larves ingéraient une quantité significativement plus élevée, avaient un taux de consommation et de croissance supérieur et augmentaient davantage leur biomasse avec des aiguilles d'épinette blanche. L'efficacité d'utilisation des aliments et le taux de croissance relatif étaient de faibles à modérément faibles pour un lépidoptère arboricole phyllophage; la vitesse d'ingestion relative était élevée. La qualité des aiguilles des trois essences d'épinettes était faible. Les aiguilles d'épinette blanche permettaient aux larves d'avoir une performance significativement plus élevée, performance mesurée à partir de plusieurs critères, alors que les aiguilles d'épinette rouge et d'épinette noire donnaient une performance égale. La supériorité des performances obtenues avec l'épinette blanche explique la plus grande vulnérabilité de cette essence à la Tordeuse des bourgeons de l'épinette.

     

Occurrence and effects of Nosema fumiferanae infections on adult spruce budworm caught above and within the forest canopy

1 Nosema fumiferanae infections in populations of both sexes of spruce budworm Choristoneura fumiferana moths, collected live above the forest canopy (canopy moths), within the tree crown (crown moths) and in drop trays (dead moths), were examined over a 5‐year period in New Brunswick, Canada. 2 The incidence of infection and of moderate–heavy infections in canopy and crown moths of both sexes increased concomitantly with moth eclosion, indicating that N. fumiferanae retards larval/pupal development, with infected moths, particularly those having higher disease loads, emerging later in the season. 3 Infection rates differed among canopy, crown, and dead female, but not male, moths. Canopy (i.e. emigrating) females had a lower incidence of infection, lower incidence of moderate–heavy infections, and had longer forewings and higher dry weights, than crown females. These results suggest that N. fumiferanae infections negatively affect aspects of female, but not male, flight performance. Regardless of infection, forewing length and dry weight of both canopy and crown females declined over the moth flight period, but infected females in both moth types were smaller than their uninfected counterparts. Forewing lengths and dry weights of moderately–heavily infected females were most severely affected. 4 Despite high annual infection rates in parents, only a small percentage of offspring (second‐instar larvae) that established feeding sites each spring were infected, indicating that high rates of horizontal transmission occurred annually throughout the larval period. 5 The present study indicates that whether N. fumiferanae infections are a debilitating sublethal factor in spruce budworm populations depends more on the disease load than on the overall incidence of infection. The potential importance of N. fumiferanae infections on various fitness parameters related to host dispersal is discussed.     

Effects of tree age and stand thinning related variations in balsam fir secondary compounds on spruce budworm Choristoneura fumiferana development,growth and food utilization

• 1 The effect of tannins and monoterpenes on the development, mortality and food utilization of spruce budworm Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) was investigated under laboratory conditions using an artificial diet. Tannins were extracted from balsam fir foliage of thinned and unthinned stands to reproduce stand thinning related variations in tannins. A mixture of synthetic monoterpenes was utilized to simulate the concentration found in young and old balsam fir trees.
• 2 Longer development time and lower pupal weight were observed for insects fed on diets with a lower nitrogen concentration and a higher tannin concentration (unthinned treatment). Tannins induced higher insect mortality at a low nitrogen concentration compared with the diet with a higher nitrogen concentration.
• 3 Approximate digestibility was higher for larvae fed on diets with high concentrations of nitrogen at both low and high concentrations of tannins. Efficiency of conversion of digested food (ECD) decreased with an increase in tannin concentration. Tannins reduced both the relative consumption and growth rate (RCR and RGR).
• 4 Monoterpenes increased spruce budworm mortality and this mortality reached almost 50% under concentrations of monoterpene typical of the young trees compared with 20% under monoterpene concentrations found in old trees.
• 5 A higher digestibility was observed for larvae fed on diet with a higher concentration of monoterpenes, whereas efficiency of conversion of ingested food (ECI), ECD, RCR, and RGR decreased with an increase in monoterpenes in the diet.
• 6 The results obtained in the present study are consistent with the defensive role of secondary compounds such as tannins and monoterpenes in the spruce budworm–balsam fir system.

     

Oviposition strategies employed by the western spruce budworm: tests of predictions from the phylogenetic constraints hypothesis

1 Predictions from the Phylogenetic Constraints Hypothesis were tested for the first time in an eruptive forest Lepidopteran species, the western spruce budworm. 2 In previous work, we established that western spruce budworm females exhibit oviposition preferences with regard to tree age, tree vigour and host species. However, there was no evidence to support a link between oviposition preference and larval performance, which supports the Phylogenetic Constraints Hypothesis. 3 Our preference data led us to test whether female budworms use oviposition strategies to select the sites where they lay their egg masses. Our experiments were designed to make direct comparisons between latent and eruptive insect herbivores with respect to two oviposition behaviours: egg retention and avoidance of conspecifics. This type of research has not previously been conducted on any eruptive forest Lepidopteran. 4 Female budworms retained eggs instead of laying them on less preferred hosts in two of three experiments, but the percentage of eggs they retained was significantly less compared to latent insect herbivores. 5 In addition, female budworms actively avoided oviposition in areas with the highest density of conspecific egg masses, but they laid egg masses in all the other locations provided. This contrasts with the pattern seen in latent insect herbivores, which consistently avoid laying their eggs near any sites already used by conspecifics. 6 Our research indicates that there are extreme differences between latent and eruptive insect herbivores with respect to egg retention and avoidance of conspecifics, thus supporting the Phylogenetic Constraints Hypothesis.     

RearingGlypta fumiferanae [Hym.: Ichneumonidae] on a multivoltine laboratory colony of the western spruce budworm (Choristoneura occidentilis) [Lep.: Tortricidae]

Methods were developed for rearingGlypta fumiferanae Viereck on a nondiapausing laboratory colony of the western spruce budworm,Choristoneura occidentalis Freeman. Both host and parasite are univoltine and undergo diapause in nature. In this study, the parasite's voltinism was synchronized with that of a nondiapausing host.G. fumiferanae mated readily in the laboratory, and 5 consecutive generations were reared with an average generation time of less than 8 weeks-much less than the 23 weeks needed for 1 generation to develop in the field. Developmental times are reported, and some aspects of behavior described.