Sampling procedures and adult sex ratios in spruce budworm

Unbiased estimates of sex ratios that reflect local abundance of adult insects are practically difficult to obtain because many gender‐specific behavioural adaptations differentially influence the catchability of males and females in commonly applied sampling procedures. Historic data on outbreak populations of spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), the major pest of conifers in Nearctic boreal forests, include dozens of sex ratio observations for 10 different sampling procedures; these data illustrate the importance of understanding the reproductive ecology of adults to contextualize sex ratio assessments. Sex ratios of resident adults (assessed by rearing field‐collected pupae to adulthood or fogging host trees with insecticide) were not different from 1:1. Sex ratios of in‐flight adults collected using Malaise traps or light traps deployed in tree canopies were consistently male‐biased, which presumably reflects the higher level of flight activity for males relative to females. Sex ratios of moths captured outside the forest canopy (presumed migrants), in contrast, were consistently female‐biased, a trend which is expected because females seeking oviposition sites are more likely to undergo migration than are males. The sex ratio among adults that died from natural causes (collected on drop trays) was not distinguishable from 1:1. In pre‐outbreak (endemic) populations, sex ratios estimated by light trapping were much more strongly male‐biased than in outbreak populations. This surprising result should, however, be interpreted with caution because little is known of reproductive ecology in endemic budworm populations.     

Polyandry across Behavioral Classes in Female Spruce Budworm

Reproduction in spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), is characterized by sedentary oviposition in the natal forest patch early in life (gravid females are incapable of flight due to their heavy abdomen) followed by windborne migrations among females that have laid >50% of their eggs; the nature of migrations (facultative or obligatory) remains unknown. The incidence of polyandry (females with more than one spermatophore in their reproductive tract) was evaluated for several thousand field-collected female budworms (data professionally archived at Canadian Forest Service) using an array of sampling procedures that can be delineated along four behavioral classes: (1) resident females collected on host trees; (2) in-flight females captured while foraging in forest canopies; (3) migrant females captured above tree canopies; and (4) females collected postmortem on drop trays. Overall, polyandry increased over time (reflecting the ageing of local populations) and with female body size (likely due to male mating preference for large, most fecund females). Polyandry did not increase postmortem realized fecundity of females. The level of polyandry was lowest among migrants, intermediate for in-flight and resident females, and highest for naturally dead females. Sedentary reproduction among young female budworms may limit gene flow and increase the risk of inbreeding; a low level of polyandry among migrants may thus be due to females not only seeking novel habitats for their progeny but also mating partners that are not closely related genetically.     

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.     

Comparison of male and female emerald ash borer (Coleoptera: Buprestidae) responses to phoebe oil and (Z)-3-hexenol lures in light green prism traps

We conducted trapping experiments for the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) in Michigan, U.S.A., and Ontario, Canada, to compare unbaited light green sticky prism traps with traps baited with phoebe oil, (Z)-3-hexenol (Z3-6:OH), or blends of other green leaf volatiles (GLVs) with Z3-6:OH. Traps were placed in the lower canopy of ash trees (Fraxinus spp.). Catches with Z3-6:OH-baited traps showed a significant male bias and these traps caught significantly more males than the unbaited controls at both sites. They were also superior to phoebe oil-baited traps and those baited with GLV blends. Catches with phoebe oil showed a significant female bias but there was no difference in the number of females captured between traps baited with phoebe oil or Z3-6:OH lures. Catches were analyzed at regular time intervals to examine the response of A. planipennis to the lures over the course of the flight season. Z3-6:OH-baited traps consistently caught more males than the controls at each interval throughout the flight season. Catches of females with Z3-6:OH and phoebe oil were significantly better than the controls early in the flight season but declined to control levels by midseason. Our results suggest that Z3-6:OH-baited green traps placed in the ash canopy would be a superior lure for detecting and monitoring A. planipennis throughout the flight season.     

Assessment of entomological indicators of Aedes aegypti (L.) from adult and egg collections in S?o Paulo, Brazil

We compared the presence and mean number of eggs from oviposition traps with the mean number of Aedes aegypti females captured by manual aspirators and by MosquiTRAPs to determine the sensitivity of each method and to standardize each indicator. The collections of adults and eggs were performed over 23 weeks in six neighborhoods of Mirassol, state of S?o Paulo, Brazil. A better assessment of indicators required larger number of MosquiTRAPs, but to quantify the number of females per house, one trap was sufficient. The sensitivities of MosquiTRAPs and manual aspirations to detect the presence of A. aegypti females were similar, but were lower compared to oviposition traps. The correlation coefficients between the number of females captured by MosquiTRAPs and manual aspirations and the number of eggs from oviposition traps were low, which may be a consequence of each method showing different stages of the mosquito life cycle.     

Reproductive Biology and Behavior of <Emphasis Type="Italic">Tranosema rostrale</Emphasis> (Hymenoptera: Ichneumonidae), a Parasitoid of Low-Density Spruce Budworm (Lepidoptera: Tortricidae) Populations

Tranosema rostrale (Brischke) (Hymenoptera: Ichneumonidae) is an important parasitoid of low-density spruce budworm Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) populations. To investigate the effectiveness of this parasitoid in attacking low-density spruce budworm populations, we conducted a detailed laboratory study on its reproductive biology and behavior including mating behavior, potential fecundity, longevity, host searching, and oviposition behavior. We found that the occurrence of mating increases with the number of males present in the cage but that it drastically decreases when females mated previously. Females may mate multiple times with different males in one breeding session and mating duration is significantly longer when a male mates the second time with the same female. Dissections of T. rostrale’s oviducts showed that it is a synovigenic species emerging with about 17 % of its lifetime egg load and develops most of its eggs in the first three days after emergence at 20 °C. Sugar, but not pollen, significantly increased the insect’s longevity compared to water. Spruce budworm larvae, silk from larvae and damaged balsam fir foliage triggered probing in T. rostrale females significantly more often than larval feces. The sequence of the parasitoid’s behaviors leading to successful attack is described as antennation, probing, insertion of the ovipositor, oviposition, and subsequent disinterest or resuming of the sequence. Defense mechanisms of the host larva such as vigorous movements, biting, and/or regurgitation and behavioral countermeasures by T. rostrale are described in detail.     

Temporal variation in abundance of male and female spruce budworms at combinatory associations of light traps and pheromone traps

A 3‐year study (2014–2016) was conducted at Rocky Harbour near the west coast of Newfoundland, Canada, to record the abundance and phenology of adult spruce budworms captured at traps, using a factorial design (light traps and pheromone traps deployed contiguously or segregated spatially). Budworms were most abundant and occurred seasonally earlier in 2014 than in 2015 and 2016; these findings held generally true for males and females. The geographic setting of Newfoundland (large island isolated from the mainland by an oceanic barrier of >100 km across) provides an ideal location to discriminate local flight from long‐range immigrations; in our study, however, immigrations cannot be ruled out for any single day of trapping due to broad overlap in emergence patterns at Rocky Harbour relative to forest stands with known populations of budworms on the mainland. Based on moderate daily variation in adult abundance, however, major immigration events (defined as external deposition of budworms with large numerical amplitude) likely did not take place at Rocky Harbor between 2014 and 2016. Males were more abundant at light traps coupled with pheromone traps, whereas abundance of males at pheromone traps was similar with or without contiguous light traps. This outcome may be mediated by lower range of attraction for light traps (usually <100 m) and (generally assumed to be several hundreds of meters). Females were equally abundant at light traps with or without pheromone traps. As expected, males were captured earlier in the season at pheromone traps than at light traps, and females occurred later in the season due to protandry. The onset of flight observed at light traps or pheromone traps in 2015 and 2016 occurred 10–15 days later than simulated predictions; caution is thus warranted as to conclusions derived on computer modeling of adult emergence.     

OVIPOSITION BEHAVIOUR AND DISPERSAL OF PERTHID A GLYPHOPA COMMON (LEPIDOPTERA: INCURVARIIDAE)

Abstract
Oviposition behaviour of the jarrah leafminer moth Perthida glyphopa was observed during the seasonal flight period in April-May. After alighting on trees, the gregarious males restricted their movements to the branches, whereas the solitary females searched for oviposition sites on the leaves. The females laid most eggs at 14–22 °C in diminishing light or overcast conditions, preferring young leaves in the lower portion of the crown. They dispersed more readily than males.     

Importance of multiple mating to female reproductive output in Diaphorina citri

Abstract The importance of multiple mating to female reproductive output in Diaphorina citri Kuwayama (Hemiptera: Psyllidae) is evaluated by grouping individual females with one or three males for 24 h (short duration) or 2 weeks (long duration) and examining oviposition over 18–19 days. For the short‐duration treatments, females lay more eggs per day when grouped with multiple males, whereas females in the long‐duration treatments lay more eggs when paired with one male. When held for 24 h with one or three males, females show a decline in fecundity beginning 10 and 15 days after mating, respectively. Total fecundity is relatively high for females paired with one male for 2 weeks, but fecundity is low and mortality high when females are held with three males for 2 weeks. In treatments in which females are held with males for 2 weeks, oviposition increases dramatically in the days after removal of males. For females paired with a male for 24 h and re‐paired for 24 h, 9 days later, fecundity remains high throughout the 18–19‐day observation period. Egg fertility does not differ among treatments, but varies over time in a manner that is similar among treatments. The present study demonstrates that, in D. citri, females require multiple matings over time to achieve high reproductive output, but oviposition is constrained by the presence of males.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

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.     

Modeling mountain pine beetle (Dendroctonus ponderosae) oviposition

Mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is a significant forest disturbance agent with a widespread distribution in western North America. Population success is influenced by temperatures that drive phenology and ultimately the adult emergence synchrony required to mass attack and kill host trees during outbreaks. In addition to lifestage‐specific developmental rates and thresholds, oviposition timing can be a source of variance in adult emergence synchrony, and is a critical aspect of mountain pine beetle phenology. Adaptation to local climates has resulted in longer generation times in southern compared to northern populations in common gardens, and the role of oviposition rate in these differences is unclear. Oviposition rates and fecundity in a northern population have been described, although data are lacking for southern populations. We assessed southern mountain pine beetle oviposition rates and fecundity in a range of temperatures using a non‐destructive technique that included frequent X‐ray imaging. We found that oviposition rate and fecundity vary independently such that a female with high oviposition rate did not necessarily have high fecundity and vice versa. Observed fecundity within the 30‐day experimental period was lowest at the lowest temperature, although estimated potential fecundity did not differ among temperatures. Females at varying temperatures have the potential to lay similar numbers of eggs, although it will take longer at lower temperatures. Southern mountain pine beetle reared in Pinus strobiformis Engelm. (Pinaceae) had a higher upper threshold for oviposition, a similar lower threshold, and slightly greater potential fecundity compared to a northern population reared in Pinus contorta Douglas. A comparison of modeled oviposition rates between the two populations, which could be influenced by host tree, suggests that differences in oviposition rate do not explain observed differences in total generation time. Our oviposition model will facilitate development of a phenology model for southern mountain pine beetle populations.     

Effect of food and temperature on the reproduction of the blowfly, <Emphasis Type="Italic">Calliphora vicina</Emphasis> R.-D. (Diptera,Calliphoridae), a popular model object in biological research

Egg maturation in Calliphora vicina is known to occur within a wide range of temperatures, from 12°C to nearly 30°C (Vinogradova, 1991). Photoperiodism has no effect on this process. Some females enter diapause already at 20°C; their fraction increases at lower temperatures and reaches 100% at 6°C. Reproducing females with eggs can survive for a long time and even lay eggs at low temperatures (4–5°C). Experiments with C. vicina from Leningrad Province revealed some effects of the diet (liver or fish) and temperature on the fly reproduction. At 20 and 25°C, 7–10-day old females begin to oviposit, but at 20°C egg maturation is observed in 98% of females feeding on liver and in only 5% of females feeding on fish. On the liver diet, the mean daily fecundity is significantly correlated with the day of oviposition but not with the temperature. At 20°C a significant correlation is observed between the mean daily fecundity and both the day of oviposition and food. The total number of eggs laid by flies after feeding on fish is half that produced after feeding on liver. The optimal conditions for Calliphora vicina cultivation are a 16-h light day, temperatures within the range from 20 to 25°C, and liver as food.     

Evaluation of tebufenozide carry-over and residual effects on spruce budworm (Lepidoptera: Tortricidae)

Laboratory and field studies investigated carry-over effects of tebufenozide on spruce budworm, Choristoneura fumiferana (Clemens). In the laboratory, third and fourth instars were fed either sublethal doses of tebufenozide (10 ppm) or water onAbies balsamea (L.) Mill. needles, reared to adulthood and allowed to oviposit on laying surfaces 1 or 14 d after being sprayed with water or tebufenozide concentrations of 17.5, 35.0, and 70.0 g/liter. Percentage adult emergence and sex ratio were not affected by larval ingestion of the tebufenozide. Also, the mean number of eggs laid on untreated wax paper by moths reared from tebufenozide-treated larvae was similar to the controls. Hence, tebufenozide did not inhibit carry-over effects on treated larvae. Oviposition on tebufenozide-treated wax paper by moths reared from untreated larvae was affected by both the substrate concentration and the age of the treatment residue. When offered treated and untreated laying surfaces simultaneously, C. fumiferana did not show a preference. However, significantly fewer eggs were laid on both laying surfaces by fewer females than when tebufenozide was absent. Residual tebufenozide on wax paper did not affect egg hatch but topical applications were toxic to eggs. Field studies appear to corroborate laboratory results and suggest that although the ingestion of tebufenozide by larval spruce budworm might not impair adult reproduction, the insecticide's presence in the environment could inhibit oviposition. This inhibition was considered to be a primary factor in tebufenozide's multi-year effects against spruce budworm populations.     

Oviposition by Trichostrongylus colubriformis (Nematoda) in vitro

Summary

Oviposition by females of the ruminant nematode Trichostrongylus colubriformis was influenced by in vitro conditions. Release of eggs declined linearly during incubation for 4.5 h. An average of 8.8 eggs was deposited by females during the first hour in vitro. Decreased pH, temperature, and osmolarity caused reduced production of eggs. Storage of females at 4°C had no effect on subsequent oviposition at 37°C. Oviposition was similar under light and dark conditions. Concomitant males or the female's age did not influence production of eggs in vitro. The release of eggs in vitro was significantly decreased by octapamine and GABA while serotonin elevated oviposition. Thus, environmental stimuli from the host may partially modulate helminth fecundity.     

Oviposition by Mamestra brassicae (L.) (Lep., Noctuidae) in relation to age, time of day and host plant

The age at which females begin to lay, the daily pattern of oviposition, the influence of host-plant material on egg-laying as well as the host preference of Mamestra brassicae L. (Lep., Noctuidae) were investigated under laboratory conditions. The females began to oviposit during the third, fourth and fifth scotophase after emergence. A great variability in daily female fecundity was observed. The highest level of oviposition occurred during the fifth day. Females oviposited during the whole scotophase, but maximal oviposition occurred during the second hour. The females without host-plant material laid fewer eggs than females with host-plant material (cabbage leaves Brassica oleracea var. capitata ). In two-choice tests, the females preferred to oviposit on cabbage rather than chrysanthemum, but there was no difference in the mean number of eggs laid on cabbage and tomato. There were significantly greater mean numbers of total eggs deposited on tomato than on chrysanthemum.     

Fecundity,longevity and caloric patterns in female Heliothis virescens: Changes with age due to flight and supplemental carbohydrate

Bomb calorimetry, survivorship and fecundity analysis of female moths indicated that feeding on 5% sucrose reduces the impact of flight activity on longevity and oviposition. Flight has only a slight to moderate effect on longevity or reproductive capacity in fed females; but in unfed females, flight significantly decreased longevity and fecundity. By comparisons to unfed or unflown females, it is concluded that female moths use lipids as the primary flight fuel prior to the initiation of oviposition. Once oviposition begins the female increases its usage of carbohydrates as the flight fuel. Older females which are unable to obtain nectar cannot maintain their flight capacity, resulting in decreased reproductive potential. Therefore, nectar quantity, quality, and likelihood of availability can be important factors determining moth longevity and reproductive and flight capacities in the field.     

Oviposition preference of Lygocoris pabulinus (Het., Miridae) in relation to plants and conspecifics

Abstract: To predict possible locations of Lygocoris pabulinus (L) in the field during the summer, we determined their oviposition preference under summer conditions. With L. pabulinus reared on potato, oviposition preference was determined for potato, tomato or green bean. As preference may depend on larval or early adult experience, the oviposition preference of bugs reared on green bean for three generations, and of bugs captured from the field 12 h prior to the experiment was also determined. All females showed a strong preference for potato plants, on which fecundity was higher. Hence, although L. pabulinus is a generalist in its feeding habits, the summer generation seems to be an oviposition specialist. Aggregation of ovipositing females does not seem to occur; similar amounts of eggs were oviposited in plants with clip cages containing conspecifics as in plants without conspecifics. More eggs were oviposited in damaged plants than in undamaged plants. Plant volatiles released upon damage may aid L. pabulinus females in finding suitable oviposition sites.     

Reduced realised fecundity in the pine looper Bupalus piniarius caused by host plant defoliation

1. Whether the potential fecundity of herbivorous insects is realised or not may depend on female behaviour, which in turn may be influenced by host plant acceptability. Female Bupalus piniarius were observed to discriminate against needles growing out the year following defoliation (current‐year needles) of its host plant Pinus sylvestris. 2. It was hypothesised that the discriminatory behaviour was due to current‐year needles being less secure as a substrate. Field and laboratory experiments were designed to test this hypothesis and to estimate the discrepancy between potential and realised fecundity when females were offered defoliated branches. 3. In a laboratory oviposition experiment, B. piniarius females were exposed to branches bearing either current‐year needles only or both mature and current‐year needles. Daily oviposition rate, egg batch size, longevity, and mature eggs and fat retained at death were recorded for each female. In field experiments, the rate at which eggs dropped from expanding needles and the capacity of neonate larvae hatching from the dropped eggs to colonise a tree were assessed. 4. Significantly fewer eggs were laid when females were exposed to defoliated branches. 5. Twenty‐six and 16% of the eggs laid on current‐year needles dropped from the needles in 1998 and 1999 respectively, whereas no eggs dropped from mature needles in 1998 and only one egg (< 1%) dropped in 1999. 6. A very small proportion of larvae hatching on the forest floor (simulated egg drop) was able to recolonise host trees. 7. These results emphasise the importance of oviposition behaviour on realised fecundity when analysing insect population dynamics. In the case of B. piniarius, egg placement, although a minor detail during the normal course of events, became of key importance when defoliation deprived females of their preferred egg attachment site.     

Effects of forced egg-retention in Aedes albopictus on adult survival and reproduction following application of DEET as an oviposition deterrent.

The insect repellent DEET (0.1% concentration), used as a mosquito oviposition deterrent in the laboratory, influenced the retention and maintenance of mature eggs by caged gravid female Aedes albopictus Skuse. This egg-retention mechanism could benefit survival because the gravid females were ultimately able to lay maintained eggs upon availability of water, but the length of forced egg-retention time reduced the number of eggs laid per female. Gravid females with retained eggs also laid a higher percentage of eggs that failed to tan, and this percentage increased with time duration of egg-retention. Percent egg hatch was not significantly affected by DEET when used as an oviposition deterrent; however, percent hatch was affected by time duration of egg-retention in both treated (exposed to DEET) and untreated (control) gravid females. The rate of egg hatch was considerably reduced after three weeks of retention; this reduction declined to zero for treated and control females at six and four weeks post-treatment, respectively. The fecundity and fertility of gravid female Ae. albopictus were affected by the time duration of forced egg-retention.