An investigation into the variations in the fine surface details of the eggs of five species of Delia (Diptera: Anthomyiidae)

The eggs of five species of Delia have been examined for the external chorionic patterning, the size and shape of the micropyle and the number, characteristics and distribution of the aeropyles using the scanning electron microscope. The species concerned are the cabbage root fly Delia radicum , the onion fly Delia antiqua , the seed corn maggot Delia platura , the wheat bulb fly Delia coarclata and the turnip fly Delia floralis.
Each species has its own clear characters and this, coupled with a significant difference in egg size, enables one species to be distinguished from another.     

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.     

Neighbouring monocultures enhance the effect of intercropping on the turnip root fly (Delia floralis)

Knowledge of insect behaviour is essential for accurately interpreting studies of diversification and to develop diversified agroecosystems that have a reliable pest‐suppressive effect. In this study, we investigated the egg‐laying behaviour of the turnip root fly, Delia floralis (Fall.) (Diptera: Anthomyiidae), in an intercrop‐monoculture system. We examined both the main effect of intercropping and the effect on oviposition in the border zone between a cabbage monoculture [Brassica oleracea L. var. capitata (Brassicaceae)] and a cabbage‐red clover intercropping system [Trifolium pratense L. (Fabaceae)]. To investigate the border‐effect, oviposition was measured along a transect from the border between the treatments to the centre of experimental plots. Intercropping reduced the total egg‐laying of D. floralis with 42% in 2003 and 55% in 2004. In 2004, it was also found that the spatial distribution of eggs within the experimental plots was affected by distance from the adjoining treatment. The difference in egg‐laying between monoculture and intercropping was most pronounced close to the border, where egg‐laying was 68% lower on intercropped plants. This difference in egg numbers decreased gradually up to a distance of 3.5 m from the border, where intercropped plants had 43% fewer eggs than the corresponding monocultured plants. The reason behind this oviposition pattern is most likely that flies in intercropped plots have a higher probability of entering the monoculture if they are close to the border than if they are in the centre of a plot. When entering the monoculture, flies can pursue their egg‐laying behaviour without being disrupted by the clover. As the final decision to land is visually stimulated, flies could also be attracted to fly from the intercropped plots into the monoculture, where host plants are more visually apparent. Visual cues could also hinder flies in a monoculture from entering an intercropped plot. Other possible patterns of insect attack due to differences in insect behaviour are discussed, as well as the practical application of the results of this study.     

Identifying sources and mechanisms of resistance in crucifers for control of cabbage maggot (Diptera: Anthomyiidae)

The cabbage maggot, Delia radicum (L.) is an important insect pest of eruciferous crops in upstate New York. This species causes considerable damage to seedlings and young plants by feeding on roots and stems, resulting in plant stand loss and yield loss. Five crucifer accessions (Brassica oleracea variety italica L.,'Green Comet'; B. oleracea L.,'Rapid Cycling' [Crucifer Genetics Cooperative 3-1 ]; B. oleracea variety botrytis L., a standard cauliflower cultivar'Amazing'; B. carinata L.; and Sinapis alba L., 'Cornell Alt 543') were evaluated to identify sources and mechanisms of resistance for D. radicum. Of the accessions tested, S. alba Cornell Alt 543 demonstrated reduced oviposition by D. radicum, reduced weights and survivorship of larvae, pupae or adults, and reduced damage to plants. Thus, S. alba Cornell Alt 543 could be a potential source for resistance to be bred into cruciferous crops for control of D. radicum.     

Binomial sampling as a cost efficient sampling method for pest management of cabbage root fly (Dipt., Anthomyiidae) in cauliflower

A binomial (presence–absence) sampling plan has been developed based on the relationship between the proportion of cauliflower plants having visible cabbage root fly eggs ( Delia radicum L.) exposed on the soil surface around the plant stem and the mean density of eggs per plant. The Kono–Sugino's model was fitted to a total of 125 population estimates, each based on 10 plant samples collected from cauliflower fields in 1994 and 1995 (P=0.001; R²=0.64). When the model was compared with an independent data set consisting of 39 population estimates collected in 1995, an analysis of covariance showed no significant differences between the regression lines. The efficiency of the binomial method was compared with absolute sampling in terms of relative precision and cost efficiency. The binomial method had a high coefficient of variation, RV ≈ 0.85, due to large biological error. In spite of this, binomial sampling was more cost efficient than the applied soil sampling when between 10 and 30 plants were examined for the presence of visible eggs.     

Comparison of the egg-laying behaviour and electrophysiological responses of Delia radicum and Delia floralis to cabbage leaf compounds

Abstract.  The behaviour and the sensitivity of adult cabbage root fly, Delia radicum and turnip root fly, Delia floralis are compared with host-plant extracts and isolated crucifer compounds previously identified as oviposition stimulants for D. radicum . The oviposition behaviour of both species is similar; 7–10-day-old females are stimulated to lay eggs by the methanol extract of cauliflower leaves that contains thia-triaza-fluorenes (CIF) as well as glucosinolates. The glucosinolate fraction is mainly composed of glucobrassicin, which alone stimulate both fly species to lay eggs. The C₅ and D_3,4 sensilla on the prothoracic tarsae of newly-emerged D. radicum contain neurones sensitive to the glucosinolate fractions tested and to glucobrassicin, whereas the CIF specifically stimulate a neurone in the C₅ sensillum. By contrast, newly-emerged D. floralis respond less to glucosinolates, especially to glucobrassicin, and have sensitive neurones to CIF in other sensilla than D. radicum . Recordings are also made from the longest sensilla present on the labellum because they are apparently sensitive to glucosinolates. By contrast to earlier investigations, no remarkable phasic-tonic responses of these neurones are seen. The two species are difficult to discriminate visually, have the same host plants, show identical host-selection behaviour, apparently respond to the same physical and chemical properties of their host-plants, but have a clearly different distribution of receptor neurones in the tarsal sensilla.     

Niche overlap and interspecific association between Chilo partellus and Chilo orichalcociliellus on the Kenya coast

The use of felt traps to estimate oviposition by the cabbage root fly, Delia radicum (L.), and turnip root fly, Delia floralis (Fall.), was compared with soil sampling at seven localities between 1992 and 1994 in Denmark and Norway. In all, 281 comparisons were made, based on 6800 samples. In 4.6% of these comparisons no eggs were found by either method. In 16% of the comparisons, 2.0±0.41 (±S.E.) eggs were found per soil sample and no eggs were found in felt traps, whereas in 0.7% of the comparisons 0.10±0.03 eggs were found per felt trap and no eggs in soil samples. When eggs were found using both methods, the ratio between soil sampling and felt traps varied from 13.1±3.2 when the egg laying rate was very low to 1.8±0.2 at high egg laying rates. Regression analysis showed significant correlation between felt trap catches and soil sampling (P<0.001), whereas locality did not influence felt trap efficiency (P=0.096). The influence of the physical conditions of felt traps, i.e., whether they were clean and dry, were analysed in a single year. Tentative action thresholds for control measures against cabbage root flies and turnip root flies were developed for cauliflower, Brassica oleracea, convar. botrytis (L.) Alef., var. botrytis, based on data from the literature and the present results.     

The effectiveness of traps used currently for monitoring populations of the cabbage root fly (Delia radicum)

Six different types of traps were compared for capturing adults of Delia radicum. Cone traps caught so few flies that it seems inadvisable for them to be used for monitoring populations of this pest. All the other traps tested were suitable for monitoring D. radicum populations. When expressed on a trap for trap basis, the large 1800 cm² Canadian traps caught most flies. When expressed as the numbers of flies caughthnit area of trapping surface, most flies were caught in the water traps. Similar numbers of D. radicum, D. platura, syrphids, blowflies and a tachinid, Eriothrix rufomaculatus were caught per unit area on each of three different vertical sticky traps. Although water traps have the advantage that they catch about 5 times as many females D. radicum/unit area as the most effective vertical sticky traps of a comparable size, their disadvantage is that they catch about 10 times as many beneficial syrphids. The reasons why water traps are so effective against D. radicum are discussed, with details of how to convert water-trap data to sticky-trap equivalents, and vice versa.     

Evaluation of entomopathogenic fungus Metarhizium anisopliae against soil-dwelling stages of cabbage maggot (Diptera: Anthomyiidae) in glasshouse and field experiments and effect of fungicides on fungal activity

The effect of two isolates of the entomopathogenic fungus Metarhizium anisopliae (Metchnikoff) Sorokin (389.93 and 392.93) on root-feeding stages of cabbage root fly, Delia radicum (L.), was studied under glasshouse and field conditions. In glasshouse studies, the effect of drenching a suspension of conidia (concentration 1 x 10(8) m(-1), 40 ml per plant, applied on four occasions) onto the base of cabbage plants infested with D. radicum eggs was compared with mixing conidial suspension into compost modules (concentration 1 x 10(8) ml(-1), 25 ml per plant) used to raise seedlings. Drench application reduced the mean number of larvae and pupae recovered per plant by up to 90%, but the compost module treatment had no statistically significant effect. Both application methods reduced the emergence of adult flies from pupae by up to 92%. Most conidia applied as a drench application remained in the top 10-cm layer of compost. Applications of the fungicides iprodione and tebuconazole, which are used routinely on brassica crops, were compatible with using M. anisopliae 389.93 against D. radicum under glasshouse conditions, even though these fungicides were inhibitory to fungal growth on SDA medium. In a field experiment, drench applications of M. anisopliae 389.93 to the base of cauliflower plants at concentrations of 1 x 10(6) to 1 x 10(8) conidia ml(-1) did not control D. radicum populations, although up to 30% of larval cadavers recovered supported sporulating mycelium. Drench applications often exhibited considerable lateral movement on the soil surface before penetrating the ground, which may have reduced the amount of inoculum in contact with D. radicum larvae.     

The relationship between cabbage root fly (Delia radicum) larval feeding and the freeze-dried matter and sugar content of Brassica roots

Five genotypes of swede (Brassica napus var. napobrassica), two genotypes of kale (B. oleracea var. acephala), and two genotypes of rape (B. napus var. napus) were each inoculated at the 8–10 true leaf stage with five cabbage root fly (Delia radicum) eggs. The percentage pupation after larval feeding on individual plant genotypes ranged from 45 to 78%, and the mean pupal weight from 6.5 to 13.0 mg. After 5 weeks, larval feeding damage had reduced root weight by up to 47%, compared with uninoculated plants. The dry matter content of undamaged roots was higher in the kales and rapes than in the swedes. Whilst the dry matter content of the rapes and swedes were not changed by D. radicum damage, that of the kales was elevated. The ethanol-soluble sugar content of the root was reduced in all cases by D. radicum larval damage. However, the effect of D. radicum damage on the concentrations of individual sugars (glucose, fructose and sucrose) was crop- and genotype-dependent. In the roots of kales and rapes, the glucose and fructose concentrations were either very low or unaffected by D. radicum damage, whilst both glucose and fructose were generally reduced in swede roots by D. radicum damage. The root sucrose concentration was either reduced or not significantly affected by D. radicum damage in all of the crop types tested. The percentage pupation and the mean pupal weight of D. radicum were inversely correlated to root freeze-dried matter content. D. radicum pupal weight was positively correlated with root fructose, glucose and ethanol-soluble sugar contents.     

Geographical origin of an introduced pest species, Delia radicum (Diptera: anthomyiidae), determined by RAPD analysis and egg micromorphology

The origin of introduction of the cabbage root fly, Delia radicum Linnaeus to the north-eastern coast of North America in the 19th century has been assumed to be from Europe. From that point of introduction, D. radicum gradually spread westward to occupy available ecological niches. DNA fingerprinting and egg micromorphology were used to determine the most likely geographical origin of the North American populations of this species. Forty-five informative RAPD loci obtained from ten primers and three criteria for egg micromorphology were studied. These characters indicated a common origin for the North American populations and a high similarity between populations from North America and north-western Europe. The results suggest a single entrance point of D. radicum into North America, probably via the north-eastern coast (New York area) from north-western Europe. The implications of this study in assisting selection of natural enemies of this important agricultural pest are discussed.     

Effect of soil temperature and moisture on survival of eggs and first-instar larvae of Delia radicum

Edaphic factors such as soil temperature and moisture influence soil-dwelling insects, whose most vulnerable stages typically are eggs and young larvae. In this study, the survival of eggs and first-instar larvae of the cabbage maggot, Delia radicum L., was measured under laboratory conditions after exposure to a range of soil temperatures and moistures. When eggs were exposed to constant temperature (20-29°C) and humidity (5-200% [wt:wt]), temperature had no significant effect on survival, whereas humidity <25% [wt:wt] caused egg mortality. The gradual exposure of eggs to high temperatures resulted in low mortality below 33°C, but <5% of eggs survived at 40°C. When first-instar larvae were exposed to constant temperature (17-29°C) and humidity (5-100% [wt:wt]), both factors as well as their interaction had a significant effect on larval survival, which was nil at 5% (wt:wt) for all temperatures but increased from 21.9 to 42.8% at 17°C and from 34.1 to 55.0% at 29°C, for soil moisture contents of 15% and 100% (wt:wt), respectively. Eggs of D. radicum are resistant to low soil moisture and high temperature conditions. Larval survival tends to increase with an increase in soil temperature and moisture. It is suggested that soil temperature be integrated into insect development simulation models instead of air temperature, to build more effective models for cabbage maggot management.     

Field evaluation of the combined deterrent and attractive effects of dimethyl disulfide on Delia radicum and its natural enemies

Delia radicum (L. 1758) is a major pest of cabbage crops in northern Europe. Due to more constraining laws relating to insecticide use, new strategies to control this pest are urgently needed. Manipulating insect behavior through infochemicals is a promising approach. The recent identification of dimethyl disulfide (DMDS) as a compound that both attracts the main predators of D. radicum and inhibits oviposition by the fly gives a challenging opportunity to develop such strategy. The aim of the present study was to confirm such potential of DMDS, in the field. Through the 8 weeks of the first egg laying peak of the fly we assessed, the potential of artificially increasing the levels of this molecule in the close vicinity of broccoli plants to 1/attract predators, 2/stimulate predatory activity and 3/limit damage done by the fly. Despite a lower number of D. radicum eggs as food resource, DMDS effectively increased predator catches in treated plots (119 Aleochara bilineata (Gyllenhal, 1810) caught in treated plot, while only 21 in control plots). However, damages done by the fly were of the same magnitude order in treated plots than in control ones. Number of D. radicum larvae and pupae recovered in plant roots were similar, despite the important decrease in eggs laid. This result, together with the observation that the numbers of eggs predated in artificial patches were lowered in the presence of the molecule, seems to indicate that increasing DMDS amounts disturbed the foraging activity of the fly predators. Consequences of these findings for the future of DMDS use in crop protection against D. radicum are discussed.     

Trap cropping effect on oviposition behaviour of the leek moth Acrolepiopsis assectella and the diamondback moth Plutella xylostella

The potential of trap cropping as a pest control strategy was explored for two specialist moth species, where oviposition behaviour determines larval distribution. Oviposition behaviour was studied at three different scales: Individual plants in the laboratory, small scale field experiments with and without cages, and medium scale field trials. The study insects were the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), a specialist on Brassica species and the leek moth, Acrolepiopsis assectella (Z.) (Lepidoptera: Acrolepiidae), a specialist on Allium species. The diamondback moth preferred Indian mustard, Brassica juncea (L.) over white cabbage, B. oleracea var alba (L.). The leek moth did not show preference when offered leek, Allium porrum (L.) and chives, A. schoenoprasum (L.). However, the leek moth did prefer to oviposit on plants of a larger biomass over smaller plants. In both the small scale and the medium scale field studies the trap crops reduced the number of eggs laid on the primary crop. The total number of eggs laid in the monoculture and the trap crop treatments were the same in the small scale field cage study, but the females distributed their eggs differently. More eggs were laid on trap crops compared with primary crops and this pattern was also validated in the medium scale field trials. In this study only the direct effects of trap cropping were considered. Indirect effects of vegetation diversity such as enhancement of natural enemies could potentially increase the efficacy of trap cropping.     

Trap cropping effect on oviposition behaviour of the leek moth Acrolepiopsis assectella and the diamondback moth Plutella xylostella

The potential of trap cropping as a pest control strategy was explored for two specialist moth species, where oviposition behaviour determines larval distribution. Oviposition behaviour was studied at three different scales: Individual plants in the laboratory, small scale field experiments with and without cages, and medium scale field trials. The study insects were the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), a specialist on Brassica species and the leek moth, Acrolepiopsis assectella (Z.) (Lepidoptera: Acrolepiidae), a specialist on Allium species. The diamondback moth preferred Indian mustard, Brassica juncea (L.) over white cabbage, B. oleracea var alba (L.). The leek moth did not show preference when offered leek, Allium porrum (L.) and chives, A. schoenoprasum (L.). However, the leek moth did prefer to oviposit on plants of a larger biomass over smaller plants. In both the small scale and the medium scale field studies the trap crops reduced the number of eggs laid on the primary crop. The total number of eggs laid in the monoculture and the trap crop treatments were the same in the small scale field cage study, but the females distributed their eggs differently. More eggs were laid on trap crops compared with primary crops and this pattern was also validated in the medium scale field trials. In this study only the direct effects of trap cropping were considered. Indirect effects of vegetation diversity such as enhancement of natural enemies could potentially increase the efficacy of trap cropping.     

Evaluating the enemies hypothesis in a clover‐cabbage intercrop: effects of generalist and specialist natural enemies on the turnip root fly (Delia floralis)

• 1 The relative importance of the resource concentration hypothesis and the enemies hypothesis was investigated for the turnip root fly Delia floralis in a cabbage–red clover intercropping system compared with a cabbage monoculture.
• 2 Delia floralis egg densities were measured as well as the activity‐densities of generalist predators in a field experiment during two growing seasons. In the second year, a study of egg predation with artificially placed eggs was conducted, in addition to a predator exclusion experiment, to estimate total predation during the season. Parasitization rates were estimated from samples of pupae.
• 3 Delia floralis oviposition was greater in the monoculture during both years. The predator activity‐densities differed between treatments and study years. The known natural enemies of Delia spp., Bembidion spp. and Aleochara bipustulata showed a strong response to a cultivation system with higher activity‐densities in the monoculture. The response, however, appeared to be caused primarily by habitat preferences and not by D. floralis egg densities.
• 4 The reduction in the number of D. floralis pupae in the intercropping may be explained by a disruption in oviposition behaviour caused by the presence of clover because neither predation, nor parasitization rates differed between cultivation systems.

     

Influence of trap surface on the numbers of insects caught in water traps in brassica crops

The contribution of the various surfaces of a water trap to the trap's overall effectiveness was studied in brassica crops by painting black different parts of fluorescent-yellow water-traps. Three pest species, Delia radicum (L.), D. platura (Meig.), and Meligethes aeneus (Fab.)/Meligethes viridescens (Fab.), together with blowflies and syrphids, were caught in large numbers. Each insect responded differently to the yellow/black traps. The numbers of insect caught indicated that the area of trap involved in capturing D. radicum was effectively twice the surface area of the water. Yellow traps for monitoring D. radicum populations could be made more selective by painting the inner wall black, as such traps caught similar numbers of D. radicum to all-yellow traps but 50% fewer D. platura, Meligethes sp. and blowflies, and 95% fewer syrphids.     

Entomophthora muscae resting spore formation in vivo in the host Delia radicum

The formation in vivo of Entomophthora muscae resting spores was investigated in the host, Delia radicum (cabbage root fly), by analysis of field data on the seasonal occurrence of E. muscae resting spores over 4 years. E. muscae resting spores in D. radicum were spherical with an average diameter of 39.4 microm, and the average numbers produced were estimated at 5.7 x 10(4) resting spores/female cadaver. Resting spores were found only after midsummer in D. radicum and almost exclusively in females. The proportion of infected females with resting spores was negatively correlated with average weekly day length after midsummer. We did not detect any significant year effect; thus, the results support the hypothesis that the photoperiod is the most important abiotic factor controlling E. muscae resting spore formation in D. radicum.     

Potential adaptation of a Q biotype whitefly population from poinsettia to field crops

Abstract The invasive Q biotype whitefly was first detected in the US on poinsettia in 2004 and is still not a pest outside of greenhouse environments in the US. To assess the potential for the establishment of the Q biotype on field crops, population cage experiments were conducted to compare the performance of a poinsettia‐derived Q population named P’06 on poinsettia and six field crops (alfalfa, tomato, melon, cotton, cowpea and cabbage). P’06 adults reared on poinsettia as nymphs laid eggs on all six field crops. Significantly more eggs were laid on alfalfa, tomato, melon and cotton than on cabbage, cowpea and poinsettia. These eggs hatched and the nymphs developed to adults on the six field crops. Relative to poinsettia, whitefly survival was similar on cowpea, alfalfa, tomato and cabbage, but significantly higher on cotton and melon. Moreover, P’06 had significantly shorter development times from egg to adult on cotton, melon, cowpea, tomato and alfalfa than they did on poinsettia. However, the F1 adults raised on the six field crops had significantly shorter lifespans and laid 11‐ to 18‐fold fewer eggs than did the F1 adults raised on poinsettia. Taken together, while P’06 may have some potential to establish on field crops, the shorter lifespans and extremely low fecundities of the F1 adults raised on the six field crops suggests that P’06 is incapable of rapidly adapting to them. Poor adaptation to field crops may explain, at least partially, why the Q biotype has not established in the US field system.     

Trap cropping to control Delia radicum populations in cruciferous crops: first results and future applications

This study evaluates the efficacy of a new approach to the control of Delia radicum populations. We suggest associating the primary crop with a trap crop that is expected to be more attractive to D. radicum females and to attract and sustain their natural enemies such as Aleochara bilineata and A. bipustulata. Various cruciferous species were compared in terms of their preference for adult D. radicum females, and performance, as estimated by larval survival. Laboratory results were complemented by field experiments in which the selected trap crops were associated with broccoli plants. The following results were obtained. Of the six different cruciferous plant species tested, Delia radicum females showed a strong preference for Chinese cabbage, with turnip also being attractive. Following the laboratory results, turnip was chosen as a trap crop because it was easier to cultivate and presented good preference and performance. In the field experiments, Aleochara adults were present in higher numbers in plots associated with turnips than in pure broccoli plots. The presence of Aleochara adults in plots with turnips improved plant protection; as fewer broccoli plants were attacked, the attacked plants were less severely damaged, and more D. radicum pupae were parasitised than in pure broccoli plots. Delia radicum females did not lay fewer eggs on broccoli plants associated with turnips. Moreover, protection and parasitism were more effective in the rows closest to the central row of turnips, suggesting that Aleochara adults limit their activity to its immediate vicinity.