Pesticide-induced mortality of natural enemies of the onion maggot,Delia antiqua [Dip.: Anthomyiidae]

Pesticide-induced differential mortality betweenDelia antiqua(MEIGEN)and several other organisms associated with its natural control [D. platura(MEIGEN),Coenosia tigrina (F.),Entomophtora muscae (COHN), andAphaereta pallipes (SAY)] were evaluated under simulated field conditions. Direct and residual differential mortalities were described for 3 herbicides (Chloro-IPC, nitrofen, and CDAA), 3 fungicides (maneb, chlorothalonil, and copper sulfate), and 1 insecticide (malathion). The recommended field application rates of these chemicals produced high positive differential mortality levels betweenD. antiqua and some of the natural mortality agents. Chloro-IPC, a preplant and midseason herbicide, induced 100% mortality ofA. pallipes over the 3-day residual test period. Modifications in behavioral orientation and death patterns ofD. antiqua adults infected withE. muscae were also noted in malathion-treated flies such that subsequent conidial dispersal was highly restricted. Michigan Agricultural Experimental Station Journal Article Number 10133.     

Biphasic effect of low temperature on completion of winter diapause in the onion maggot,Delia antiqua

The effect of low temperature on completion of winter diapause was investigated in the onion maggot, Delia antiqua (Diptera: Anthomyiidae). Diapause was completed under constant diapause-inducing conditions of 15 degrees C and 12L-12D, without any exposure to lower temperature. The pupal period for 50% adult emergence was 117 days. None of the cold treatments at 5.6 degrees C examined in the present study advanced adult emergence; on the contrary, they delayed it. Detailed analyses of the results revealed that diapause development in D. antiqua comprises two phases which differ in sensitivity to low temperature, with the phase shift occurring at around day 60 at 15 degrees C and 12L-12D. In the first phase of diapause development, low temperature (5.6 degrees C) had no effect on diapause development. In the latter phase, by contrast, diapause development was retarded in proportion to the duration of cold treatment.     

Laboratory transmission and in vivo incubation of Entomophthora muscae (Entomophthorales: Entomophthoracae) in the onion fly,Delia antiqua (Diptera: Anthomyiidae)

A high-moisture infection chamber was used for the in vivo transmission of Entomophthora muscae within laboratory populations of the onion fly, Delia antiqua. This cadaver-to-fly transmission procedure provided an average experimental infection rate well above 95%. Laboratory infection and temperature-dependent incubation rates of E. muscae were further examined in adult populations of D. antiqua. The time from initial exposure until host death and pathogen sporulation was accurately predicted using a second-order function of the incubation temperature. A developmental base temperature of approximately 5°C was estimated, with 105 degree-days being the average number of heat units required between host infection and death. E. muscae transmission between D. antiqua and D. platura, two insect pests typically associated with Michigan onion production, was verified under laboratory conditions.     

Modification of female onion fly,Delia antiqua (Meigen), reproductive behavior by male paragonial gland extracts (Diptera: Anthomyiidae)

Egg depositional rates of onion flies, Delia antiqua(Meigen), injected thoracically with extracts of male paragonial glands were identical (14.5 eggs/female/ day) to those of normally mated females. Moreover, when continuously exposed to males, extract-injected females refused to mate and produced unfertilized eggs for the duration of the > 15- day experiment. For this normally monocoitic dipteran, <1 male equiv of paragonial secretion completely reproduced the ovipositional responses characteristic of normal mating, and this effect required no involvement of the genitalia or genital chamber. We suggest that the receptor for the active chemical (s) (sex peptide?) would be an excellent target for biorational insect control by sterilization. Moreover, these primer sex pheromones might play an important role in insect reproductive isolation and evolution.     

Delaying onion planting to control onion maggot (Diptera: Anthomyiidae): efficacy and underlying mechanisms

Onion maggot, Delia antiqua (Meigen) (Diptera: Anthomyiidae), is an important pest of onion, Allium cepa L., in northern temperate areas, especially in the Great Lakes region of North America Management of D. antiqua relies on insecticide use at planting, but insecticide resistance can cause control failures that threaten the long-term viability of this strategy. Delaying the time onions are planted was investigated as an alternative management approach for D. antiqua and the ecological and behavioral mechanisms underlying host age and insect relationships were examined in laboratory and field experiments. Delaying onion planting by two to four weeks reduced damage to onions by 35 and 90%, respectively. Onions planted later emerged later and this reduced the period overwintered flies had to oviposit on the plants. Moreover, flies tended to lay few to no eggs on these young, late-planted onions. As anticipated, D. antiqua laid 4-8 times more eggs on older onions than on young onions, and older onions were more resilient to injury caused by D. antiqua neonates compared with younger onions. However, the resiliency to maggot attack lessened as the density of D. antiqua increased from 2 to 10 eggs per plant, which probably explains why greater levels of maggot damage are typically observed in early onion plantings compared with later plantings. Delaying onion planting until mid-May reduced D. antiqua damage without jeopardizing the period required to produce marketable yield, but this cultural tactic must be combined with other management strategies to prevent economic loss.     

Purification and immunological characterization of superoxide dismutase of the onion maggot,Delia antiqua

Cytosolic superoxide dismutase (SOD) of the onion maggot, Delia antiqua, was purified to apparent homogeneity by ammonium sulfate fractionation followed by anion exchange, hydrophobic interaction, and gel filtration chromatographies. Native molecular mass was estimated as 32,000 daltons. SDS-PAGE revealed only one subunit of 16,000 daltons, indicating that SOD is a homodimer. Isoelectric focusing revealed 3 charge isomers of pls 5.3, 5.5, and 5.7. The specific activity of purified SOD was 4,250 U/mg protein. A monoclonal antibody (MAb, aSOD2B7) raised against Delia SOD recognized only SOD of the same genus, but another MAb (aSOD1H11) recognized SOD of Drosophila melanogaster as well. © 1995 Wiley-Liss, Inc.     

Cyromazine seed treatments to control onion maggot, Delia antiqua, on green onions

Cyromazine seed treatments were evaluated for onion maggot control in green onion crops. The more tolerant to organophosphates of two populations of onion maggots was chosen for further research, based on the results of adult assays in a Potter spray tower. In the laboratory, first-instar mortality was compared between film-coated seed treatment and soil drench treatment. The LC50 for the film-coated seed treatment was approximately one fourth that of the soil drench treatment. In choice assays, no significant difference was observed between the number of eggs deposited on seedlings grown from film-coated seeds with cyromazine and film-coated seeds without cyromazine. Field studies demonstrated that all rates of cyromazine seed treatment resulted in protection of onion plants from onion maggot damage. Green onions may not require as high a rate of cyromazine for control of onion maggots as the rate established for use in dry onions for two reasons: the seeding rate is much higher for green onions resulting in more AI/ha for a given amount of AI/kg of seed, and a given percentage of stand loss is more difficult to detect in green onions than in dry onions.     

Onion maggot (Diptera: Anthomyiidae) resistance to chlorpyrifos in New York onion fields

A larval immersion bioassay was developed to identify susceptibility of onion maggot, Delia antiqua (Meigen) (Diptera: Anthomyiidae), to chlorpyrifos and to determine whether this assay could be used to predict control in onion fields. Laboratory colonies were established from larvae collected in New York onion fields during 2003 and 2004, providing us with test insects to use in bioassays. The larval assay effectively determined susceptibility of D. antiqua to chlorpyrifos, and results were congruent with an adult bioassay. However, use of similar-aged larvae (4 d old) in the assays was critical because larvae became more tolerant to chlorpyrifos as they aged. In a field survey, six of the 13 populations had LC50 values above the recommended field rate of 3,600 ppm (range 4,031-6,869). Over two successive seasons in the same field, susceptibility of D. antiqua to chlorpyrifos decreased in two of three fields (by 45 and 42%) and remained the same in another field, indicating that resistance is not predictable from year to year. Based on the relationship between damage in the field and LC50 values from 11 of the populations mentioned above, all five populations that had LC50 values above the field rate caused unacceptable levels of damage, whereas five of six populations that had LC50 values below the field rate did not cause serious damage.     

Nonwoven fiber barriers for control of cabbage maggot and onion maggot (Diptera: Anthomyiidae).

We investigated the use of nonwoven fiber barriers for control of cabbage maggot, Delia radicum (L.), and onion maggot, D. antiqua (Meigen). The barriers consist of arrangements of minute fibers loosely intertwined in "web" form. Results from a greenhouse experiment showed that manually applied graphite fibers placed at the base of broccoli plants reduced the number of D. radicum eggs by 64-98%, and that efficacy increased with greater fiber density. Using a melt extrusion process, we devised a method for on-site creation of nonwoven fibers of ethylene vinyl acetate (EVA). In field trials with broccoli and onion plants, EVA fibers significantly reduced the number of cabbage and onion maggots infesting plants. Fiber barriers provided comparable control to standard insecticide applications. The addition of blue, yellow, red, or black pigments, as well as optical brighteners that absorb UV light did not enhance fiber efficacy. Incorporation of capsaicin olfactory repellent to EVA also did not enhance fiber efficacy. Nonwoven fiber barriers may offer an alternative to insecticides for control of cabbage maggot and onion maggot and possibly other insect pests. Additional research is needed to improve the application process and to identify economically feasible and biodegradable compounds for fibers.     

The relationship of age and ovarian development to mating of Delia antiqua (Diptera: Anthomyiidae)

Abstract. The relationship between age and sexual receptivity for male and female onion flies, Delia antiqua (Meigen) (Diptera: Anthomyiidae), and factors affecting ovarian development were examined under various conditions. Confinement in small arenas had a significant negative effect on the rate of oogenesis over the first 10 days of adult life and, in some cases, on the percentage of females inseminated. The rate of oogenesis was not affected by male population density, but high density of females in large arenas appeared to have a priming effect. Few males in single-pair mating bioassays conducted over 24 h were sexually mature at 3–4 days post-eclosion, but > 50% mated when aged 6–7 days. Although females first mated at 3–4 days post-eclosion when confined for 24 h with sexually mature males, most females mated at 6–7 days of age when oocytes in the terminal ovariole position comprised 50% of total egg volume (stage 7 or greater on a 10-stage scale). No females aged 3 days were mated in no-choice and two-choice bioassays with sexually mature males over 24 h. Previtellogenic females (stage 3) were not inseminated, and ovarian development was only correlated weakly ( r = 0.48) with mating. Frequency of mating in mixed-sex groups of twenty flies was comparable with that in single pairs. However, more females were inseminated at 3–4 days, probably as a result of multiple matings by a few precocious males. These results do not support the hypothesis that females mate only when ovarioles are mature.     

Identity and bioactivity of oviposition deterrents in pine oil for the onion maggot, Delia antiqua

The oviposition deterrent properties of pine oil (Norpine 65, Northwest Petrochemicals, Anacortes, Washington) for the onion maggot, Delia antiqua (Meigen), were verified using a two-choice bioassay with onion oil as an attractive control. The principal deterrent property of this pine oil was found to reside in three monoterpenes, 3-carene, limonene, and p-cymene, which were the primary constituents identified in the most deterrent of two fractions made by preparative gas chromatography of steam-distilled pine oil. At release rates of 220, 320 and 320 g per 24 h in two-choice bioassays these monoterpenes respectively caused 73.2, 65.4 and 56.3% deterrency of oviposition, while the ternary mixture released at 320 g per 24 h caused 88.6% deterrency. The ternary mixture also caused 62.5% deterrency in a no-choice bioassay. Of eight other monoterpenes tested, myrcene, -phellandrene, -terpinene, -phellandrene, -terpinene, terpinolene, and -pinene were significantly deterrent in declining order, while -pinene was inactive. The ternary mixture was released from glass capillary tubes or flexible plastic cylinders in further bioassays that challenged caged females to oviposit around the base of 35 potted onion seedlings with release devices placed on the soil surface. The most effective deterrency (85.3%) was achieved at a release rate of 280 g per 24 h per pot if plastic cylinder devices were deployed 24 h before the treated pot was exposed to D. antiqua females. If female D. antiqua were given only a treated pot, deterrency of oviposition on potted onion seedlings was significant, but low (11.7–63.2%). Because of incomplete efficacy, a monoterpene-based deterrent formulation would be best used operationally if combined with other deterrents, or if it were integrated with some other tactic.     

Laboratory evaluation of entomopathogenic fungi against larvae and adults of onion maggot (Diptera: Anthomyiidae)

Laboratory experiments were done to measure the susceptibility of larvae and adults of the onion maggot, Delia antiqua (Meigen) (Diptera: Muscidae: Anthomyiidae) to 27 isolates of entomopathogenic fungi from four genera [Beauveria Vuillemin, Lecanicillium (Petch) Zare & W. Gams, Metarhizium Sorokin, and Paecilomyces Bainier]. A novel bioassay was developed for D. antiqua larvae by using a diet based on mixed vegetable powder. When evaluated in a virulence screen, the fungal isolates caused less mortality of D. antiqua larvae than adults. Only three isolates caused > 50% mortality of larvae, whereas 12 isolates caused > 50% mortality of adults. Fungal species was a statistically significant factor affecting the mortality of larvae but not of adults. The fungal isolates causing the most mortality of larvae tended to belong to Metarhizium anisopliae (Metschnikoff) Sorokin. Two M. anisopliae isolates (389.93 and 392.93) were evaluated in dose-response bioassays. The median lethal concentrations of the isolates against larvae were 6.1 x 10(7) conidia ml(-1) for isolate 389.93 and 7.6 x 10(7) conidia ml(-1) for isolate 392.93. The emergence of adult flies from pupae was reduced at high concentrations of conidia (3.0 x 10(8) and 1.0 x 10(8) conidia ml(-1)). The median lethal concentrations of the isolates against adults were 1.7 x 10(7) and 4.0 x 10(7) conidia ml(-1), respectively. Some of the fungal isolates examined may have potential as biological control agents of larvae of D. antiqua and related species.     

Crop sequence effects on root maggot (Diptera: Anthomyiidae: Delia spp.) infestations in canola

Strong market demand for canola, Brassica napus L., has prompted some western Canadian producers to increase the frequency of this crop in rotations with other crop species, but the impact of this practice on canola insect pests has not been determined. Here, we investigate 12 cropping sequences involving canola over a 3-yr period (2008-2010 inclusive) at five locations across western Canada. Cropping sequences varied from continuous production of two herbicide-tolerant canola varieties, to production in two of 3 yr, to canola production in one of the 3 yr. Treatments analyzed were the frequency and timing of canola within the rotational sequence. Damage by larvae of root maggots (Diptera: Anthomyiidae: Delia spp.) to canola taproots increased as the study progressed, particularly in 2010 after canola had been grown continuously for 3 yr. Yield declined with continuous canola production, and differences were greatest in 2010. At mean canola crop prices for 2010, the yield reduction from continuous production amounted to economic losses of approximately Can$282-$377/ha. Crop quality, in terms of oil and protein concentrations of harvested seed, was affected more by crop variety than cropping sequence. Crop sequence effects for root maggot damage, yield, and seed quality were relatively stable in the presence of environmental (location) variation. Results of our study suggest that continuous canola production could be unsustainable over the long-term even though market forces currently provide incentive for this practice.     

Chemical control of onion fly, Delia antiqua

In nine trials done in 1972-4 seed treatments reduced attacks of onion fly larvae very effectively and most of the organophosphorus formulations gave protection up to harvest, 12–17 wk after sowing. Pirimiphos-ethyl and isofen-phos each at 10 or 20 g a.i./kg seed controlled onion fly larvae well and they were not seriously phytotoxic in any of the trials. Chlorpyrifos, trichloronate, ethion, iodofenphos, chlorfenvinphos wettable powder or triazophos all at iog a.i./kg seed were also effective, but they were tested in fewer trials. Diazinon at 10 or 20 g a.i./kg seed, carbophenothion and chlorfenvinphos liquid seed dressing at 10 g a.i./kg seed controlled onion fly well, but in some trials they were phytotoxic; methiocarb, mecarphon and oxamyl were much less effective. At 1–12 kg a.i./ha chlorfenvinphos granules applied over the rows shortly before the attack was expected was the best granular formulation, but it was not as effective as the better seed treatments.     

Upregulation of a desaturase is associated with the enhancement of cold hardiness in the onion maggot, Delia antiqua

Cold-acclimated non-diapause pupae, and summer- and winter-diapause pupae of the onion maggot, Delia antiqua (Diptera: Anthomyiidae), show marked cold hardiness as compared with intact non-diapause pupae. Homeoviscous adaptation of cellular membranes is crucial to enhance the cold hardiness of organisms, and Delta9-acyl-CoA desaturases have been assumed, albeit without experimental evidence in insects, to play a key role in the adaptation. We cloned the cDNA of a desaturase gene (Dadesat) from D. antiqua, which is most likely to encode Delta9-acyl-CoA desaturase. Expression of Dadesat mRNA in the brain, midgut, and Malpighian tubules of cold-acclimated and diapause pupae was upregulated 2-10 fold, correlating well with the increase in cold hardiness. In the pupae with enhanced cold hardiness, palmitoleic and oleic acids, the presumed products of Dadesat, in the phospholipids were significantly increased. These findings suggest that the increase in the expression of Dadesat contributes to enhanced cold hardiness in D. antiqua through the production of these unsaturated fatty acids.     

Comparison of the circadian eclosion rhythm between non-diapause and diapause pupae in the onion fly,Delia antiqua

The influence of pupal diapause on adult eclosion rhythm of Delia antiqua was investigated. When non-diapause and diapause pupae were exposed to various photoperiods at 15, 20 and 25 °C, both of them emerged as adults close to the light-on time, but the phase of eclosion varied with photoperiod and temperature. Moreover, there was a significant difference in the eclosion time between non-diapause and diapause pupae; the eclosion peak of diapause pupae was earlier than that of non-diapause pupae. When non-diapause and diapause pupae were transferred to constant darkness (DD) after having experienced LD 12:12 at 15, 20 and 25 °C, both showed circadian rhythmicity in eclosion. Although the free-running period (τ) decreased slightly as temperature increased in both non-diapause and diapause pupae, the latter tended to show shorter τ than the former. This observation suggests that the observed difference in eclosion time in LD cycles between non-diapause and diapause pupae is due to differences in τ.     

Variation due to maternal age in the onion root maggot,Hylemya antiqua (Meigen)

A variety of characteristics were studied in populations of the onion root maggot, Hylemya antiqua, to determine the extent to which the age of the female producing a batch of eggs might affect heterogeneity in her offspring. Early-born offspring were found to be reproductively most successful. They had the longest mean expectation of life, a relatively low mortality rate into mid-life, the highest average fecundity, the highest mean rate of egg production, and they produced the highest percentage of female offspring. On the other hand, mid-born offspring were hardiest. They also had a long mean expectation of life, comparable to that for early-born offspring but, in addition, they had the lowest sustained mortality rate and the greatest ability to survive food stress. Their robustness may have been acquired at the expense of certain reproductive capabilities however, in that their average fecundity was lower than their early-born siblings', and their rate of egg production was more variable. Late-born offspring were the most mortality-prone and the least fecund, but they had the shortest mean generation time. These differences must be evaluated in terms of their effects on the rate of natural increase and on population growth. Maternal age was found to influence pupal size, although the provenance groups differed significantly from each other and, in fact, showed opposite trends within their respective cohort groups. Size itself, and the ability of a species to alter the size at which it pupates, may be characteristics which have evolved to fit a specific set of environmental conditions.     

Onion fly,Delia antiqua,oviposition and mating as influenced by insect age and dosage of male reproductive tract extract (Diptera: Anthomyiidae)

One hundred percent of virgin female onion flies,Delia antiqua, receiving 1/20 of a male equivalent of an aqueous extract of mature male reproductive tract remained unmated in the presence of males and began laying unfertilized eggs at a normally mated rate of about 20 eggs/female/day. The 50% behavioral response (BR₅₀) fell between 1/40 and 1/20 of a male equivalent. Sex peptide responses are not always all-or-none. Some females receiving extract at 1/40 male equivalent oviposited at an intermediate rate. Moreover, at low sex peptide dosages, some females were fully activated ovipositionally but were receiptive to mating. A low level of sex peptide was present in 1-day-old males. Sex peptide titer rose with age until plateauing by 6 days posteclosion. Males began mating at 3 days, when they first had ample mature sperm; 50% of 6-day-old males mated. The mean number of females inseminated per male exposed to an excess of virgin females over 24 h was 4.3±0.6 (±SE). Presence of mature eggs was not always a prerequisite for mating, although probability of insemination was correlated with egg maturation. One-day-old preovipositional females receiving 1/20 of a male equivalent of extract began ovipositing when they had mature eggs at 5–6 days old. Therefore, sex peptide may act early and permanently or have a long half-life and affect behaviors once females reach sexual maturity. Male flies provide females with an excess of sex peptide in many cases.D. antiqua males transferred ca. 5–10 times more sex peptide than necessary to activate females fully. We suggest this excess is related to the speed of female response. It is yet unclear whether sex peptide potency or titer in Diptera has become exaggerated by intra- or intersexual selection.