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.     

远东地区影响针叶树天然更新的球果种实害虫———球果花蝇 (双翅目 :花蝇科)( 英文)

球果花蝇是远东地区的重要球果种实害虫。尤其是在我国东北地区 ,由于球果花蝇的危害已严重影响了落叶松种子产量 ,进而影响到落叶松的造林与更新。本文系统地总结了近些年危害落叶松、云杉、冷杉种实的球果花蝇的研究进展及存在的问题。文中还涵括了作者的一些原始研究数据和尚未发表的新结果。     

Characteristics of summer diapause in the onion maggot,Delia antiqua (Diptera: Anthomyiidae)

Characteristics of summer diapause in the onion maggot, Delia antiqua, were clarified by laboratory experiments. Temperature was the primary factor for the induction of summer diapause in this species. The critical temperature for diapause induction was approximately 24 degrees C, regardless of the photoperiod. At 23 degrees C, the development of the diapausing pupae was arrested the day after pupariation, when about 7% of the total pupal development had occurred in terms of total effective temperature (degree-days). The most sensitive period for temperature with regard to diapause induction was estimated to be between pupariation and "pupation" (i.e., evagination of the head in cyclorrhaphous flies). Completion of diapause occurred at a wide range of temperatures (4-25 degrees C): The optimal temperature was approximately 16 degrees C, at which temperature only five days were required for diapause completion. The characteristics of summer diapause in D. antiqua are discussed in comparison with those of summer dormancy in a congener D. radicum and those of winter diapause in D. antiqua.     

Choices and Consequences of Oviposition by Hylemya (Delia) Sp. (Diptera: Anthomyiidae)

Hylema sp.⁵females oviposit on the undersides of sepals of developing buds of both Ipomopsis aggregata and Polemonium foliosissimum. Eggs deposited on the latter are significantly more likely to be fully protected by the sepal than are eggs deposited on the former. Unexposed eggs have a significantly greater likelihood of successfully developing to the larval stage than do exposed eggs. The difference in frequency of egg exposure on the two plant species can be attributed to differences in sepal morphology: I. aggregata sepals are significantly narrower than those of P. foliosissimum. The hypothesis that females preferentially oviposit on larger flowers was unconfirmed by a manipulative choice experiment. Plants differing in the size of their flowers were potted together and presented to Hylemya in arrays in the field. Flowers of the larger-flowered pair were no more likely to be oviposited on than flowers of the smaller-flowered pair. However, there were significant negative correlations between the corolla length and the percentage of flowers laid on per day at each of two sites. There was also a significant positive correlation between the corolla width and the percentage of flowers laid on at one site. Thus females appear to be using some measure of flower morphology, or a correlated trait, in making oviposition decisions. The degree to which Hylemya is making suboptimal choices between host plant species is discussed and requires further examination.     

Specificity of accessory gland extracts. in three Delia fly species (Diptera: Anthomyiidae)

Abstract. Extracts of testes and male accessory (paragonial) glands made from three species of Delia (onion fly {D.antiqua) , seedcorn fly (D.platura) , and cabbage fly (D.radicum)) were injected into conspecific virgin females. Extracts of paragonial glands, but not testes, from onion, seedcorn and cabbage fly males stimulated oviposition and suppressed mating when injected into conspecific virgin females. When extracts of paragonial glands from males of these species were injected into heterospecific virgin females, the extracts of D.antiqua and D.platura were fully cross-reactive with respect to oviposition; interspecific injection stimulated oviposition at the level of the conspecific mated controls. Injection of D.radicum extract fully activated the D.antiqua and D.platura ovipositional response. D.antiqua extract caused mating inhibition and partial oviposition in D.radicum; that of D.platura had no effect on either oviposition or mating inhibition in D.radicum. These results suggest that D.antiqua and D.platura are more closely related to one another than either is to D.radicum , and agree with published anatomically-based phylogenies and a genetic distance calculation based on eight enzyme loci. The occurrence of sex peptide cross-reactivity, though asymmetrical, between D.radicum versus D.antiqua and D.platura indicates that, functionally, sex peptides have changed little during the evolution of this genus. An emerging pattern of broad cross-reactivity within genera suggests that sex peptides are not an initiator of reproductive isolation.     

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.     

Cold resistance and overwintering survival of the cabbage root fly,Delia radicum (Anthomyiidae), and its parasitoid,Trybliographa rapae (Cynipidae), in England

Summary Overwintering Delia radicum (L.) in a field of swedes (Brassica napus L.) near Ascot, Berks., England, were exposed to soil temperatures below 10°C on 176 days from 21 October 1983 to 22 April 1984, but no temperatures below 0°C were recorded. Collections of D. radicum taken at monthly intervals from 1 November 1983 to 30 April 1984 showed that parasitism by the cynipid Trybliographa rapae (Westw.) and by the staphylinid Aleochara bilineata Gyll. was the main source of mortality. A substantial increase in parasitism by A. bilineata occurred during November, but much of the increase was by superparasitism of pupae previously parasitized by T. rapae. Mortality from causes other than parasitism was greater in the November and December collections (c. 22%) and in the spring (c. 12%) than during the winter (c. 3%) and could not be attributed to low temperatures.In all collections, most of the adult D. radicum (c. 90%) emerged within 230°D_5.6. The mean number of °D_5.6 to eclosion did not change from 1 November to 5 March but decreased significantly by 2 April. Late-emerging adults remired 259 to 992°D_5.6 to eclosion and no changes in the nean number of °D occurred among collections.Among unparasitized D. radicum, individual supercooling points showed a strong peak at c.-23°C. A significant proportion of pupae with supercooling points above-20°C were found only in the 1 November 1983 and the 30 April 1984 collections. The high supercooling points in the November collection may have comprised apparently healthy but moribund individuals, whereas in the April collection they may have included individuals that had initiated postdiapause development. The mean supercooling point of individuals in the low category (supercooling points 20°C) did not vary among monthly samples. Parasitism by T. rapae increased the variability in supercooling points, resulting in a few individuals with lower, and many more with higher supercooling points than among unparasitized individuals. The mean supercooling point increasing from the collections of November and December to those of January to April.Puparia containing unparasitized D. radicum pupae were heavier and contained more water than those with parasitized pupae, but neither group showed significant ranges over winter. Supercooling points were positively correlated with puparial live weight among unparasitized but not among parasitized pupae. Supercooling points were not correlated with water content for either group.Parasitism did not affect the occurrence or concentration of sugars and polyhydric alcohols (all<1% of fresh weight), and trehalose, glucose and mannitol were the most abundant. D. radicum can be considered to be over-protected from lethal freezing in the pupal stage and its high supercooling capacity in England may persist because it is conferred by the structural properties of the dipteran puparium and pupa within it and therefore is not subject to selection pressures.     

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.     

Orientation behavior of newly hatched larvae of the cabbage maggot,Delia radicum (L.) (Diptera: Anthomyiidae), to volatile plant metabolites

Newly hatched larvae of the cabbage maggot, Delia radicum(L.), responded to various olfactory stimuli from their host plants and oriented themselves by the concentration gradients of these stimuli. Allyl and ethyl isothiocyanate (NCS), characteristic metabolites of host Brassicaceae, elicited positive taxis, but at a higher concentrations were repellent. Benzyl NCS was neutral; only at the highest concentration was it slightly repellent. Phenyl, cyclohexyl, and butyl NCS were repellent in the highest amounts used. Volatile compounds emanating prevailingly from surface parts of plants were both attractive (hexanol, hexanal, cis-3-hexen-1-ol, linalool) and repellent (hexylacetate, cis-3-hexenylacetate, benzaldehyde, myrcene, terpinene, -pinene, limonene). While the attractiveness was associated with NCS group and with unsaturated and saturated alcohols and aldehydes with an optimal sixcarbon chain length, the repellency was probably caused by a cyclized carbon chain and an acetate group. The activity of a compound was strongly influenced by its concentration.     

Effects of seeding date and canola species on seedling damage by flea beetles in three ecoregions

Flea beetles ( Phyllotreta striolata (Fabricius), Phyllotreta cruciferae (Goeze) and Psylliodes punctulata Melsheimer) (Coleoptera: Chrysomelidae) are chronic threats to canola seedling establishment throughout the Northern Great Plains of North America. We conducted field studies in AB, Canada, from 2001 to 2004 at Lethbridge, Lacombe and Beaverlodge in the southern, central and northern regions of the province, respectively, to assess the impacts of seeding date (fall, April, May), and canola species ( Brassica rapa L., Brassica napus L.) on flea beetle damage to canola. Flea beetle damage was generally low at Lacombe but reached economic threshold levels at other sites. Although our results were not consistent in all years, seeding date had opposite effects on flea beetle damage to canola seedlings at the southern and northern sites. In the south, both canola species planted in April escaped flea beetle damage unlike the May-seeded plots, which required insecticide spraying in 2003. The numbers of flea beetles on sticky cards sampled during the seedling stage of canola were also lower in early-planted plots than in those planted in May. In the north, flea beetle damage was lower in the May-planted plots compared with those planted earlier. To maximize yields, canola growers need to plant as early as possible in all agro-ecoregions; our study showed that crops in the central and northern regions will be at greater risk of flea beetle damage than in southern regions.     

Comparison of nectar use and preference in the parasitoid Trybliographa rapae (Hymenoptera: Figitidae) and its host,the cabbage root fly,Delia radicum (Diptera: Anthomyiidae)

This study investigated differences in flower preferences between the parasitoid Trybliographa rapae Westwood (Hymenoptera: Figitidae) and its host, the economically important pest of cruciferous crops, the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The data obtained were used to suggest selective food plants in conservation biological control programmes for control of D. radicum. The attraction of both insect species to floral odours emitted from nine different plant species, their ability to access nectar from four of these species and the effect of the most promising plant species on insect longevity were determined. Naive T. rapae females were significantly attracted to flower odours from Fagopyrum esculentum Moench. (Polygonaceae) and repelled by Coriandrum sativum L. (Apiaceae) and Borago officinalis L. (Boraginaceae). In addition, T. rapae gained weight when exposed to F. esculentum, Anethum graveolens L. (Apiaceae) and Lobularia maritima Desv. (Brassicaceae). In contrast, naive D. radicum females showed attraction to most of the flowers. The longevity of both T. rapae and D. radicum increased significantly when they were provided with flowering A. graveolens and F. esculentum. In addition to the laboratory studies, a semi-field experiment was made to study the impact of flowering F. esculentum on the ability of T. rapae to parasitise D. radicum larvae. Significantly more larvae were parasitised in cages where a floral resource was present. The findings are discussed in the context of a Brassica agroecosystem.     

The spatial distribution of the wheat-bulb fly,Leptohylemyia coarctata (fall.) (Diptera: Anthomyiidae)

The spatial distribution of the eggs, larvae, pupae and adults of the wheat-bulb fly was investigated by fitting 42 sets of data comprising 1334 samples to the Poisson and negative binomial distributions, and by using the power law (S²=am^b). In general, the tests indicated that all stages were aggregated and fitted the negative binomial model.     

Female meiosis in the onion fly and cabbage root fly (Diptera: Anthomyiidae).

The sequence of female meiosis was investigated in two populations of the cabbage root fly (Delia radicum) and three populations of the onion fly (D. antiqua). In contrast with the completely achiasmate males, both species showed high levels of recombination in females. However, significant differences in chiasma frequency occurred between individuals within populations and between the populations. It was not uncommon to find aneuploidy of the X chromosomes. The autosomes occasionally showed asynapsis or desynapsis, but normal disjunction of univalents was facilitated by distance pairing.     

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.     

Attempts to control cabbage root flies Delia radicum L. and Delia floralis (Fall.) (Dipt., Anthomyiidae) with entomopathogenic fungi: laboratory and greenhouse tests

One laboratory and three greenhouse experiments were conducted to study the pathogenicity and efficacy of Finnish isolates of entomopathogenic hyphemycetous fungi against cabbage root flies. In Petri dishes, exposure to 1.5 × 10¹⁰ spores of Metarhizium anisopliae per dish caused 40–50% mortality of undifferentiated second- and third-stage larvae of Delia floralis , and 1 × 10⁷ spores per dish caused 40–50% mortality of Delia radicum larvae. In one greenhouse test, 1 × 10⁸ and 1 × 10⁹ spores of M. anisopliae and Paecilomyces fumosoroseus reduced the root damage of head cabbage by 20–70% compared with untreated controls, although this was not accompanied by significant reductions in the number of pupae. Only M. anisopliae consistently grew out of larvae and pupae of D. floralis during incubation that followed their recovery from the soil at the end of an experiment testing different formulations of M. anisopliae and Beauveria bassiana , but the frequency of the latent infections of the pest by M. anisopliae was not associated with reduced severity of damage to seedlings of head cabbage.     

Competition between larvae of wheat bulb fly, Delia coarctata (Anthomyiidae), for winter wheat shoots in an insectary

Newly hatched wheat bulb fly larvae were added to 32 wheat seedlings with one shoot and three to four leaves, in large pots in an outdoor insectary so that the numbers of available shoots per larva were 32, 16, 8, 2, 1 and 1/2. Competition for shoots between larvae, measured by the number of shoots attacked and the emergence of flies, occurred only when there was 1 or 1/2 shoot/larva. Fewer flies also emerged from shoots when the number available was one or less than one shoot/larva. In the field, egg numbers in autumn equal to or less than 5×10⁶/ha (2×10⁶/acre), suggest fewer than the expected carrying capacity of winter wheat growing at the usual plant population except in spots where the density is greater than average because of aggregation.Résumé Les larves de Delia coarctata attaquent les pousses du blé d'hiver en janvier et février. Une larve se nourrit sur une pousse et peut se déplacer sur une autre pour achever son alimentation avant pupaison. Le dispositif expérimental comporte 32 jeunes plants de blé d'hiver, comportant chacun une pousse et 3 à 4 feuilles, dans des pots installés dans un insectarium en plein air; des larves venant d'éclore sont réparties en nombre tel que la compétition varie en parallèle avec le nombre de pousses disponibles par larve: 32, 16, 8, 2, 1 ou 1/2 pousse par larve. La compétition mesurée d'après le nombre de pousses attaquées ne se manifeste que s'il y a seulement 1 ou 1/2 pousse par larve. Peu de mouches éclosent quand le nombre de pousses dont dispose chaque larve est égal ou inférieur à `.Dans les champs, le nombre d'oeufs à l'automne est égal ou inférieur à 5×10⁶/ha, ce qui est en dessous de la capacité d'accueil du blé d'hiver à l'hiver à l'égard de la population de ce parasite, excepté en certains points où le nombre d'oeufs et donc le nombre de larves est plus élevé qu'en moyenne, par suite de cette aggrégation.     

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.     

Effects of high-temperature stress and heat shock on two root maggots, Bradysia odoriphaga and Bradysia difformis (Diptera: Sciaridae)

Bradysia odoriphaga and B. difformis (Diptera: Sciaridae) are devastating pests of vegetables, ornamentals and edible mushrooms. In Chinese chive fields, the two Bradysia species occur with similar regularities: outbreaks in spring and autumn, and population decreases in summer. Temperature may be an important factor restricting their population abundance in summer. Here, we performed a life-table study under constant high temperatures and assessed the tolerance of two Bradysia species to heat shock. Life parameters of the Bradysia species indicated slow developmental rates, and low survival rates and fecundity, when the temperature was higher than 30 °C. At 34 °C, individuals were unable to reach the adult stages from eggs. Moreover, temperatures above 36 °C showed lethal effects, decreasing their survival rates. The median lethal time (LT50) values of 4th instar B. odoriphaga and B. difformis larvae were 46.82 and 32.97 h, respectively, while the values at 38 °C were 2.12 and 1.51 h, respectively. The 4th instar larvae and pupae possessed higher thermotolerance levels than adults and eggs, indicating sensitivities to heat stress. Moreover, B. odoriphaga was more thermotolerant than B. difformis. Thus, weak thermotolerance levels may restrict their occurrences during the period of summer heat, and the difference in thermotolerance levels between the two species may be related to their regional distributions.     

Fungus gnats, Bradysia spp. (Diptera: Sciaridae), and other arthropods in commercial bagged soilless growing media and rooted plant plugs

Fungus gnats, Bradysia spp., in greenhouses cause economic losses to horticultural producers by damaging young root systems during plant propagation, by spreading soilborne diseases, and by reducing the marketability of the crop. In a greenhouse cage study, our observations suggested that bagged soilless growing media or rooted plant plugs from wholesale distributors may be sources for the introduction of fungus gnats into commercial greenhouse facilities. To evaluate these possibilities, carefully collected samples of bagged soilless growing media stored in the greenhouse, as well as bagged soilless growing media and rooted plant plugs delivered from midwestern wholesale distributors, were incubated under controlled conditions in the laboratory. Fungus gnats emerged from soilless media stored in the greenhouse, soilless media delivered from wholesale distributors, and from rooted plant plugs delivered from wholesale distributors. These results demonstrate that pasteurization of even bagged soilless media may be essential to effectively managing greenhouse populations of fungus gnats. However, pasteurization is not an option for responding to contamination of rooted plant plugs. Preliminary evidence is provided that application of entomopathogenic nematodes may offer potential as a method for managing fungus gnats in plant plugs, so long as treatment is early. Other arthropods found contaminating soilless media and rooted plant plugs included the western flower thrips, Frankliniella occidentalis (Pergande), Collembola, Acari, Formicidae, Staphylinidae, Psychodidae, and other Diptera.