Influence of epicuticular waxes from white spruce and balsam fir on feeding behavior of the eastern spruce budworm

Fourth-instar larvae of the eastern spruce bud-worm Choristoneura fumiferanaClem. (Lepidoptera: Tortricidae) were shown in two-choice feeding tests to respond differently to epicuticular waxes from different white spruce and balsam fir trees. Larvae also showed different preferences for various wax fractions obtained by separation on silicic acid.Deceased.     

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.     

Oviposition Behavior of the Eastern Spruce Budworm Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae)

We documented 13 behaviors associated with oviposition in unfertilized and fertilized spruce budworm moths, Choristoneura fumiferana, by videotaping active moths with a macro lens. Apart from resting, the most pronounced behaviors were probing, drumming + probing, and egg laying. Probing was a bending of the abdomen and extension of the ovipositor to touch the substrate in a rocking, back and forth motion. Drumming + probing involved tapping of the substrate with the pro- and mesothocacic legs with concurrent bending of the abdomen as above. Egg laying was the actual process of egg deposition onto the needle surface of a balsam fir twig. Both the frequency and the duration of these behavioral elements varied depending on the mating status of the female. Unfertilized females exhibited a higher frequency of probing and egg laying. The duration of probing was longer in unfertilized females, while drumming + probing and egg laying were longer for fertilized females. This study is the basis for future work on the chemosensilla associated with the perception of host-plant surface chemicals by ovipositing females.     

Disruption of male spruce budworm orientation to female moths by pheromone and pheromone analogues in a wind tunnel

Abstract. The ability of male spruce budworm moths, Choristoneura fumiferana (Clem.), to orientate to and locate virgin female moths was investigated in a laboratory wind tunnel. When presented with two caged ‘target female’ moths, more than 99% of the male moths locked on and flew directly to the females. When confined for 3 h before release, downwind of a three-dimensional array of thirty-five evenly distributed female moths (a density of 40/m³), over 90% of the males still successfully oriented to one or another of the females, although not always to the target females. With an array of thirty-five septa containing either E-11-tetradecenyl acetate on E-11-tetradecenol there was no visible change in behaviour; >90% of the males flew directly to the two target females, which indicates that neither the acetate nor the alcohol has potential for mating disruption. An array of thirty-five septa containing a 955 blend of E- and Z-11-tetradecenal, the main components of the spruce budworm sex pheromone, significantly reduced the numbers of male moths able to locate the target females, and effects were concentration-dependent. The highest concentration, which gave a release rate similar to that of a calling female, produced the same level of disruption as the thirty-five females, but sensory fatigue became a factor in addition to ‘false trail following’. Lower concentrations were significantly less effective, with no evidence of sensory fatigue. At all concentrations significant numbers of males flew to the septa (false trail following), but when male moths were left uncaged in the tunnel for a further 3h, most eventually located the females. This suggests that false trail following alone will not prevent mating. The level of disruption was not improved by the addition of 2% tetradecanal to the 955 blend. Similar levels of disruption to the 955 blend were obtained with an 80:20 blend. A 5050 blend still caused a significant amount of false trail following, but less than the other blends. The ability of male spruce budworm to orientate to off-blend trails was enhanced if the males were first exposed to the natural, female-produced pheromone.     

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.     

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.     

Responses of the western spruce budworm to varying levels of nitrogen and terpenes

Summary An agar diet study using western spruce budworm populations from Idaho and New Mexico was carried out to determine the effects of varying concentrations of nitrogen, beta-pinene, and bornyl acetate on larval growth and survival. Increased availability of nitrogen resulted in increased larval growth rate and survival to the adult stage. Larval growth rates from the Idaho population were higher on the high-nitrogen diet than were growth rates from the New Mexico population. The high level of beta-pinene improved larval growth at the high-nitrogen concentration. Bornyl acetate significantly reduced larval growth at both the low and high levels of nitrogen although the effect was greatest with the high-nitrogen diet. High bornyl acetate concentrations also significantly reduced survival to the adult stage. At high-nitrogen levels, a high concentration of bornyl acetate reduced larval growth rates and adult survival to a level similar to that occurring at the low nitrogen and low bornyl acetate concentrations.     

The influence of understory nectar resources on parasitism of the spruce budworm Choristoneura fumiferana in the field

1 Nectar provided by flowering plants has been suggested as an important factor enhancing parasitism rates of herbivorous insects. Artificial nectar has been shown to increase parasitoid longevity and fecundity in laboratory studies. 2 We studied the influence of understory nectar on parasitism of the spruce budworm Choristoneura fumiferana in a field experiment in which we either removed understory flowering herbs or sprayed a sucrose solution on the understory vegetation in 0.25 ha quadrats. 3 Cohorts of laboratory-reared second instars were transferred the field to evaluate parasitism in the quadrats. The larvae were harvested as fifth instars and reared for parasitoids. 4 Parasitoids of early instars caused greater mortality in the supplemental sucrose treatment than in the vegetation-removal treatment or the control. Parasitoids of older larvae did not respond to the treatments. Larger-scale treatments may be necessary to elicit a response by the larger parasitoids that attack older larvae.     

Misleading correlations: The case of the Canada warbler and spruce budworm

In a recently published article Sleep et al. ( 2009 ) suggested that 30-yr declines in Canada warbler (1975–2005) based on Breeding Bird Survey (BBS) data could be attributed to 30-yr declines in spruce budworm through a series of analyses based principally on correlations. We demonstrate that the relationship does not hold when a longer-term data set (1968–2008) is used for the analysis. Sleep et al. ( 2009 ) also demonstrated a positive relationship between correlations of Canada warbler abundance and time and spruce budworm defoliation and time using provincial data sets. We examined the underlying BBS data and found that there were insufficient observations of Canada warbler in the western provinces (Alberta and Manitoba) to support the conclusion because most Canada warbler observations occurred far (>100 km) from any budworm defoliation. As well, we used a density-dependent stochastic population growth model as proposed by Sleep et al. ( 2009 ) and found only 2 significant relationships (Nova Scotia, Alberta) between Canada warbler population growth rate and budworm defoliation at the provincial scale. We conclude that little analytical evidence exists to support the idea that Canada warbler decline is a function of spruce budworm decline. © 2011 The Wildlife Society.     

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.     

The fine-scale population dynamics of spruce budworm: survival of early instars related to forest condition

Abstract.  1. A lagged, density-dependent relationship between survival of early instars and host-tree condition is revealed during outbreaks of spruce budworm, Choristoneura fumiferana Clem. Persistent damage to hosts leads to deterioration of the stand.
2. Resource limitation affects survival during early-instar dispersal of spruce budworm. Impediments to distinguishing these events with estimates of survival were overcome with a simple model that describes the dispersal and survival processes. The model was used to analyse a recent 15-year population series from Black Sturgeon Lake and two historical datasets from Green River, in Canada.
3. Defoliation-induced damage to the trees resulted in increased losses of spring-emerging larvae that are dispersing in search of feeding sites. Losses were further exacerbated by biotic factors such as maternal fecundity, rates of infection by the pathogen, Nosema fumiferanae , and by weather-related effects on the foraging period.
4. Survival of early-stage budworm larvae in persistent outbreaks declined and the likelihood of other density-related factors such as rate of mortality from natural enemies increased. These results may reconcile outstanding differences in interpretation of the role of the forest resource in spruce budworm population dynamics and point to a common process linking the dynamics of other well-known budworm species.     

Response of the western spruce budworm to temperature and dose of a virus, a growth regulator, and an organophosphate

Pathogenicity of nuclear polyhedrosis virus (NPV) to western spruce budworm larvae, Choristoneura occidentalis Freeman, increased as temperature increased. In constrast, at temperature extremes, mortality increased for a specified dose of malathion or diflubenzuron, while at the optimal temperatures (20–25°C), mortality decreased. Malathion was the fastest acting insecticide, followed by diflubenzuron and then NPV.Simulation results considered temperature dependent developmental rates of the larvae, energy consumption, time to achieve total mortality, and mortality at specified doses of insecticides. Low temperatures provided the greatest reduction in the amount of energy consumed by spruce budworm relative to the energy consumed when no treatment was applied. This was due to both the percent mortality and because total mortality was realized in the larval stage while at the higher temperatures, mortality was not fully realized until the pupal stage.

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Résumé La température modifie la toxicité ou la pathogénie du virus de la polyhédrose nucléaire (NPV), du diflubenzuron, et du malathion consommés par Choristoneura occidentalis Freeman. La pathogénie du NPV augmente avec la température, si bien qu'il y a une relation linéaire entre mortalité et température. Pour le diflubenzuron et le malathion, la température est liée à la mortalité d'une façon gradiatique. Aux températures extrêmes, la mortalité augmente pour une dose donnée de malathion ou de diflubenzuron, tandis qu'aux températures optimales (20 à 25°C), la mortalité diminue. Des modèles multiples ont été utilisés pour évaluer les effets ci-dessus.Le moment de la mortalité finale observée et la mortalité cumulative ont été modifiés par la température, mais non par la dose d'insecticide. L'effet le plus rapide est obtenu avec le malathion, suivi par le diflubenzuron et enfin le NPV.Selon les résultats de simulation,—tenant compte des taux de développement larvaires à différentes températures, de la consommation d'énergie, du temps nécessaire pour obtenir la mortalité totale calculée et de la mortalité calculée, pour une dose donnée d'insecticide à une température particulière —, ce sont les basses températures qui réduisent le plus l'énergie consommée par la population par comparaison avec l'énergie consommée en l'absence de traitement. Ceci est dû au pourcentage de mortalité calculée et au fait que la mortalité est toujours obtenue au stade larvaire, tandis qu'aux températures plus élevées, la mortalité se poursuit jusqu'au stade nymphal.

     

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.     

Chemoreception of amino acids by female fourth- and sixth-instar larvae of the spruce budworm

An electrophysiological approach was used to record the responses of maxillary styloconic sensilla of fourth- and sixth-instar larvae of the spruce budworm Choristoneura fumiferana to 14 amino acids. One cell in the lateral styloconic sensillum was identified as an amino acid-sensitive neuron. All of the amino acids tested, except l-proline and l-arginine, were detected by this cell. Arginine did not evoke a response from either the medial or lateral styloconic sensilla. Proline evoked responses from a cell in the medial styloconic sensillum. It is known from previous behavioural work that l-proline is a phagostimulant and l-valine inhibits feeding in Choristoneura; we thus further characterized the responses to these two amino acids. For both instars, l-proline was detected as low as 0.001 mmol/l and the maximal response was at 50 mmol/l. Stimulation of fourth- and sixth-instar larvae with l-valine showed that the maximum firing frequency was obtained at 1 mmol/l. Above and below this concentration, firing frequency decreases. Sensory responses to the amino acids stimuli did not correlate with known behavioral responses to similar stimuli.     

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

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

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.

     

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.     

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.     

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.     

Oviposition preference of western flower thrips for cucumber leaves from different positions along the plant stem

While the distribution of herbivorous insects over leaves along the stem often shows a peak at some distance from the apex this does not necessarily reflect an innate preference as alternative explanations can be provided such as impact of predators and inter- or intraspecific competitors. It is of interest to determine which factors shape the distribution of insects over the leaves of a plant. Do leaves from different positions differ in suitability for insects and is that reflected in the insect's preference, or are other factors involved? In this paper we assess how the herbivorous insect western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), chooses among leaves from different positions relative to the apex of cucumber, Cucumis sativus (L.) plants. On leaf discs of a susceptible and three partially resistant cucumber accessions, thrips reproduction was highest on apical leaves and lowest on basal leaves. In dual-choice essays thrips females preferred younger leaves over older leaves for oviposition in all cucumber accessions tested, as was predicted from the no-choice assay. This indicates that differences in leaf suitability are an important factor in determining thrips distribution on cucumber plants.