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.     

Larval dispersal from potential hosts within a population of a generalist herbivore, Choristoneura rosaceana

The dispersal behavior of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) first instar larvae was studied in the laboratory. The objectives were to investigate the proximal factors influencing larval dispersal and to establish whether a correspondence exists between larval host acceptance and performance. A dispersal bioassay was validated by demonstrating the presence of a positive correlation between larval host acceptance in the laboratory and in the field. Larval age and family origin, as well as host species attributes were shown to influence larval dispersal rates. Seasonal changes in host plants slightly changed the rank order of larval host acceptance. Leaf texture and the availability of refuges on host plants seemed to be important factors influencing the rate of larval dispersal. Plant odor appeared to be used by the larvae to locate leaves. Nitrogen content of plant species corresponded to larval dispersal rates, but the cause of this association is unclear. Larval dispersal did not match host suitabilities as measured by larval performance. The relationship between host preference and suitability in the obliquebanded leafroller is discussed in an ecological and evolutionary perspective.     

Developmental polymorphism: a major factor for understanding sublethal effects of Bacillus thuringiensis

Developmental polymorphism in Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) was induced by Bacillus thuringiensis (Berliner) in the laboratory. The B.t. formulation concentration induced a quadratic response on the incidence of developmental polymorphism. A double application of Bacillus thuringiensis at fourth and fifth instar respectively resulted in a higher occurrence of developmental polymorphism than a single application at fifth instar. Smaller larval weight prior to B.t. exposure resulted in higher incidence of developmental polymorphism. However, larvae exhibiting developmental polymorphism exhibited larger pupae and an extended larval period. Thus, developmental polymorphism and Bacillus thuringiensis ingestion together prolonged the development time but induced opposite effects on pupal weight. These findings suggest that the calculation of sublethal effects on pupal weight and development time that include developmental polymorphism as an uncontrolled source of variation could lead to an inadequate understanding of insect response in any stress-induced experiments.     

Infectivity of Strongyloides venezuelensis is influenced by variations in temperature and time of culture

The present research investigated the influence of temperature and time of larvae culture on the infectivity of Strongyloides venezuelensis. Mice were infected s.c. with 1500 larvae of S. venezuelensis maintained at 28 °C for three days of culture (dc), 28 °C for seven dc or 18 °C for seven dc. On days 1, 3, 5, 7, 14 and 21 post-infection the animals were sacrificed and cell numbers in the blood, peritoneal cavity fluid (PCF), broncoalveolar fluid (BALF), cytokines, immunoglobulins, number of parasites and eggs/g of feces were quantified. Results demonstrated an increase in eosinophils and mononuclear cells in the blood, PCF and BALF of infected mice. Larvae at 28 °C/3dc induced earlier eosinophils in the PCF and BALF as opposed to larvae at 28 °C/7dc and 18 °C/7dc. Larvae at 28 °C/7dc induced higher synthesis of IL-4, IL-5 and IL-10 on days 5 and 7 post-infection. Larvae at 28 °C/3dc in culture induced higher synthesis of IL-12 than larvae of seven dc, but time in culture induced better synthesis of IFN-γ after larval migration had ceased and only adult worms were present. Larvae at 28 °C/3dc in culture induced higher synthesis of IgG and IgG1 and expelled less female parasites than larvae cultivated for seven days. In conclusion, it was observed that the infectivity of S. venezuelensis is influenced by variations in temperature and time of culture.     

Host plant exploitation within a population of a generalist herbivore, Choristoneura rosaceana

Feeding performance of larvae of the obliquebanded leafroller, Choristoneura rosaceana (Harris), was estimated on apple and three alternative hosts in the Okanagan Valley. The obliquebanded leafroller is native to the study area and apple was introduced in the valley at the beginning of the century. Components of insect fitness were measured to help formulate predictions about the evolution of host preference in this generalist herbivore. In the summer, females attained a higher pupal weight on apple than on trembling aspen, wild rose and snowberry. Males performed best on apple and trembling aspen. In the field, females arising from caterpillars fed on apple leaves had a 26% greater mean fecundity than if fed on alternative hosts. The diet also affected development time; females initiated pupation later on trembling aspen than on the other hosts. Finally, no differences were found in larval survival nor in the proportion of larvae diapausing when fed on different hosts. These components of fitness indicate apple as the most suitable host in the summer.In the fall, diet influenced larval propensity to initiate diapause in the second or third instar, the time elapsed from hatching to diapause initiation and the weight of the instars initiating diapause. However, no difference in overwintering survival among larvae fed on different hosts was detected. The rank order of water and nitrogen contents of leaves of the host plants did not match the rank order of larval feeding performance in either season. A high suitability of apple and its abundance in the Okanagan valley are factors that would favor the evolution of an increased utilization of this host by the obliquebanded leafroller.     

Toxicity of insecticides to obliquebanded leafroller, Choristoneura rosaceana, larvae and adults exposed previously to neem seed oil

Adult and sixth instar obliquebanded leafroller, Choristoneura rosaceana (Harris), exposed previously as fifth instar larvae to sub-lethal concentrations of neem, Azadirachta indica A. Juss., seed oil contained in artificial diet were more susceptible to topically applied pyrethroid, carbamate and organophosphate insecticides than C. rosaceana reared on control diets. Increased susceptibility to insecticides did not result from reduced vigour, as measured by reductions in adult or larval weights, but is instead correlated with previously demonstrated reductions in detoxication enzyme activities. The modest increases in susceptibilities, not exceeding ca. 4.5-fold, for C. rosaceana exposed to neem were achieved with subjects from a susceptible laboratory colony; a larger response could be expected for resistant field populations having elevated detoxication enzyme levels. Neem-based insecticides could be useful tools for the management of orchard pests that have developed resistance to synthetic neurotoxins. These findings also contribute to a better understanding of the numerous physiological changes that occur in larval Lepidoptera following exposure to neem extracts.     

Assessment of three techniques for measuring natural enemy inflicted mortality of leafroller larvae in commercial orchards

Measuring natural enemy (NE) inflicted mortality presents a serious challenge in managed habitats, such as apple orchards, where the crop-plant is frequently treated with a wide range of insecticides. Toxic residues keep natural populations of pests at too low levels to sample and prevent the use of techniques in which laboratory reared insects are released directly on foliage and used to measure NE activity. Three methods to implant obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) larvae were tested: (1) larvae placed on potted apple trees, (2) larvae placed on excised leaves inserted in florists’ waterpicks and (3) larvae placed on branches placed into water filled Polyvinyl chloride (PVC) pipes. Use of potted trees allowed accurate estimation of NE inflicted mortality; however, this method was very labor intensive. The waterpick method required less labor, but it underestimated larval mortality caused by NEs. The use of PVC pipes to hold infested branches combined the benefits of the other two methods tested. This technique was simple to use and provided accurate estimates of NE inflicted mortality.     

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.     

Investigations of Mechanisms of Pheromone Communication Disruption of Choristoneura rosaceana (Harris) in a Wind Tunnel

Although atmospheric treatment with synthetic pheromone is used commercially to control several lepidopteran pests, little is known about how mate-finding behavior is altered by this procedure. Mechanisms of disruption of the mate-finding behavior of Choristoneura rosaceana (Lepidoptera: Tortricidae) were examined in wind tunnel bioassays. Male moths were exposed to synthetic sex pheromone or pheromone components prior to or during upwind flight to a calling virgin female. The proportion of males successfully contacting a calling female was reduced only when the synthetic pheromone treatment was presented simultaneously. A synthetic source of the attractive four-component pheromone placed upwind of a calling female was more effective than a less attractive two-component blend in achieving disorientation. Habituation of the central nervous system does not appear to be a significant factor in disruption of mate-finding behavior of male C. rosaceana, as exposure to pheromone prior to the bioassay did not alter the proportion of males that achieved subsequent upwind flight. Disruption of mate-finding behavior of C. rosaceana in a wind tunnel is probably the result of a combination of mechanisms including adaptation of antennal receptors, camouflage of the female-produced plume, and false-trail following which contributed to the additive disruption effect observed with the most attractive four-component pheromone.     

Prevalence of Nosema sp. (Microsporidia: Nosematidae) during an outbreak of the jack pine budworm in Ontario

Microsporidia are believed to play little or no role in outbreaks of the jack pine budworm, Choristoneura pinuspinus Freeman (Lepidoptrera: Tortricidae), because the short duration (2–4 years) of those outbreaks may not permit significant build-up of the pathogen. We conducted the first survey of Nosema sp. (Microsporidia: Nosematidae) over the course of a recent jack pine budworm outbreak in Ontario. Between 2004 and 2010 the outbreak defoliated a cumulative total of 1.78 million ha. Microscopic examination of ∼15,000 overwintering larvae collected over 6 years in sites with densities of 3 larvae per branch or more revealed widespread occurrence of Nosema at generally high infection intensities. The pathogen was present in 69.5% of the 518 plots that were monitored. Prevalence of infection was generally low (below 40% in 84% of plots with infected larvae) but reached high levels (80–95%) locally and increased rapidly in most infestations within 1–2 years of onset. We hypothesize that the habit of early-instar larvae to feed on developing male flowers (pollen cones) after spring emergence is critical in allowing rapid build-up of Nosema by increasing efficiency of horizontal transmission (higher density of both infected larvae and egested spores). Nosema infection may contribute to the complexity of jack pine budworm outbreak patterns by affecting egg recruitment and early-instar survival at the stand level in concert with known effects of budworm-induced reductions in pollen cone production on those processes.     

Feeding habits of larval Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret (Israel) II. Experimental study

Interactions between the larvae of Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret, Israel, and their zooplankton prey were studied experimentally. Prey species preference and size selectivities were measured. Larvae were hatched in the lab from eggs collected in the field, and fed different food items in various concentrations. The food items included lake zooplankton, algae, and commercial pellets. It was shown that small, first feeding larvae (7–8.5 mm SL) prefer small bodied zooplankters (< 180 µ). The effect of these food sources on larval growth was measured. It was found that larval Mirogrex grew at a higher rate when fed zooplankton prey sized from 63 µ–250 µ. Food items smaller than 63 µ, larger than 250 µ and Scenedesmus sp., produced less than optimal growth rates. The importance of Mirogrex feeding habits and their potential influence on the Kinneret ecosystem is considered.     

Recycling of entomopathogenic nematodes in Delia radicum and in other insects from cruciferous crops

Larvae of Deliaradicum Linnaeus (Diptera: Anthomyiidae) wereexposed to Steinernema feltiae Filipjev(Rhabditida) to study the ability of thenematodes to fulfill their life cycle (recycle)in this host. In addition, the recycling ofSteinernema and Heterorhabditis(Rhabditida) species was studied in larvae ofthe following insects: Meligethes spp.Stephens (Coleoptera: Nitidulidae),Dasyneura brassicae Winnertz (Diptera:Cecidomyidae), Ceutorrhynchus assimilisPaykull, C. pallidactylus Marsham (Coleoptera: Curculionidae) and Mamestrabrassicae Linnaeus (Lepidoptera: Noctuidae).All larval instars of D. radicum werestudied (larvae live and pupate in soil)whereas the final instars of the remaininginsects were studied (larvae only pupate in soil).On average, the following numbers of infectivejuveniles were produced in the different hosts:1,200–1,400 in C. assimilis; 700–1,300 inMeligethes spp. and 47,000 in M.brassicae. In D. radicum, around 400infective juveniles were produced on average inthe smallest larvae and around 3,500 in thelargest larvae. The highest number of nematodesrecorded in a D. radicum larva (thirdinstar) was 9,500 infective juveniles. Thenumber of nematodes produced in D.radicum cadavers was positively related to thesize of the insect host, but large variationwas observed.     

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.     

Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.     

Does aggregation behavior of codling moth larvae,Cydia pomonella,increase the risk of parasitism by Mastrus ridibundus?

Mastrus ridibundus is a specialist hymenopteran parasitoid that parasitizes last-instar larvae or prepupae of the codling moth, Cydia pomonella. Foraging females eavesdrop on an aggregation pheromone produced by cocooning larvae. We investigated whether larvae that cocoon in aggregation experience a greater rate of parasitism than larvae that cocoon in isolation. In wind tunnel experiments, 10 larvae in aggregations were more readily located by female M. ridibundus than 10 larvae well separated from each other. Similarly, aggregations of 30 larvae were more attractive to female M. ridibundus than those of 3 larvae. In cage experiments, larval cocooning in aggregation or isolation had no effect on the mean rate of parasitism and the mean number of eggs deposited per parasitized host. In Petri-dish experiments, the location of larvae within an aggregation significantly affected their rate of parasitism, with those in the center of an aggregation completely shielded from parasitism. Our data suggest that aggregation behavior by C. pomonella larvae does not appear to increase the rate of parasitism. The increased risk of aggregated larvae to be detected by M. ridibundus is likely offset by diluted parasitism risk and structural refugia effects that larvae in aggregation experience. As an egg-limited parasitoid, female M. ridibundus can parasitize on average only one larva in an aggregation, with the likelihood of parasitism for each larva being inversely proportional to the number of larvae in that aggregation.     

The foraging locus: behavioral tests for normal muscle movement in rover and sitter Drosophila melanogaster larvae

We used Drosophila melanogaster larvae with different alleles at the foraging (for) locus in a variety of behavioral tests to evaluate normal muscle usage of rover and sitter phenotypes. The results show that sitter and lethal sitter alleles of for do not affect larval behavior through a mutation which affects larval muscle usage. In general the behavior of rovers and sitters differed on food but not on non-nutritive substrates. Rovers and sitters moved equally well on non-nutritive substrates, and measures such as the time to roll over and length of forward stride showed no significant strain differences. Larvae with different alleles at for did not differ in body length. Rovers took more strides, not longer ones, than sitters while on foraging substrates. We conclude that differences in larval locomotion during foraging found in larvae with different alleles at for can not be explained on the basis of muscle usage alone. It is more likely that for affects larval ability to perceive or respond to the foraging environment.     

Effect of dietary selenium supplementation on resistance to baculovirus infection

We have examined the effects of dietary selenium (Se) supplementation on larval growth and immunocompetence of the lepidopteran pest, the cabbage looper, Trichoplusia ni. Supplementation of the diet of T. ni larvae with 10–20 ppm Se resulted in a 1 day delay in pupation. The effects of the addition and/or removal of dietary Se on total Se bioaccumulation and sequestration were determined by neutron activation analysis of pupae. Early penultimate instar larvae moved from selenium containing diet to basal diet lost total pupal Se content down to the level of those fed basal diet. Conversely, larvae moved from basal diet to diet containing additional Se rapidly attained pupal Se levels comparable to larvae fed Se throughout larval development. Therefore, dietary Se is rapidly accumulated or lost during larval development, but significant amounts are sequestered from diet into pupae. Larvae were reared on diet supplemented with 5 or 10 ppm Se until the onset of the penultimate instar then infected per os with increasing concentrations of the fatal baculovirus Autographa californica multiple nucleopolyhedrovirus (AcMNPV). Larvae fed Se in the penultimate and ultimate instars were more resistant to viral infection than larvae not fed Se in the final instars. This study indicates that dietary Se levels rapidly impact Se assimilation and sequestration and that tissue Se levels are an important factor in resistance to AcMNPV infection in larval T. ni.     

Life cycle and growth ofTrichostegia minor (Curtis) in temporary woodland pools (Trichoptera: Phryganeidae)

Larvae of the caddisTrichostegia minor (Curtis) were collected from four woodland pools in The Netherlands, three of which are temporary, from August 1986 till June 1987. Eggs and larvae of this species proved to be very well adapted to drought, freezing, strongly fluctuating pH and alkalinity levels and prolonged oxygen deficit. The life cycle ofT.minor in a small woodland marsh overgrown byCalla palustris took one year. Adult flight period started at the end of May. Oviposition took place independent of water. Hatching of the eggs started in September and was probably induced by immersion. During the larval stage from September until May, 5 instars could be distinguished by the size of the head capsule. Growth of instars I, II and III during autumn was moderate. Most larvae overwintered as instar III or IV. Possibly there was a larval diapause during winter. In spring rapid growth to instar V took place prior to pupation. Growth rate, expressed as the increase of mean individual dry weight was highest from March to April (2.05±0.75% DW.m^–2.d^–1). In extremely shallow water growth in spring was initially more rapid compared to growth in deeper water. During winter the growth rate decreased to 0.038±0.071% DW.m^–2.d^–1. Net annual production based on the changes of momentary biomass was 183.2±31.7 mg DW.m^–2.y^–1 or 177.2±31.3 mg AFDW.m^–2.y^–1. Production loss during the winter season was 75.1±10.8 mg DW.m^–2.y^–1 or 72.3±10.6 mg AFDW.m^–2.y^–1.     

Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Factors affecting oral regurgitation by larval spruce budworm

This study evaluated factors that influence the regurgitation behaviour of sixth instar spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), reared on balsam fir, Abies balsamea (L.) Mill. (Pinaceae), under various experimental conditions in the laboratory. Upon physical disturbance, larvae discharged a median volume of regurgitant of 0.4 μl when fed and 1.6 μl when food‐deprived. Larvae deprived of food for 24 or 48 h disgorged more regurgitant than larvae feeding on balsam fir foliage, and the effect was consistent for laboratory‐reared and field‐collected larvae. The water content of the foliage fed upon by larvae had no immediate impact on the volume of regurgitant; following a 24‐h period of food deprivation, however, larvae that previously fed on fresh foliage discharged >2.5 times more regurgitant than larvae that previously fed on dry foliage. Self‐regulated regurgitation by larvae, measured using the number of regurgitant stains on filter paper, was >10 times higher when larvae had access to balsam fir foliage than when they were starved. The number of larvae confined inside the Petri dish (one or four individuals) had a relatively small effect on regurgitation. Larvae were deterred from feeding when balsam fir needles were entirely covered with regurgitant, but not when only a portion of the foliage was treated. These results suggest that the regurgitant does not serve as resource marking or spacing pheromone. The high level of regurgitation by larvae after contact with ants suggests that the regurgitant has evolved in part as a defence mechanism against natural enemies.