Companion planting – do aromatic plants disrupt host‐plant finding by the cabbage root fly and the onion fly more effectively than non‐aromatic plants?

Brassica and Allium host‐plants were each surrounded by four non‐host plants to determine how background plants affected host‐plant finding by the cabbage root fly (Delia radicum L.) and the onion fly [Delia antiqua (Meig.)] (Diptera: Anthomyiidae), respectively. The 24 non‐host plants tested in field‐cage experiments included garden ‘bedding’ plants, weeds, aromatic plants, companion plants, and one vegetable plant. Of the 20 non‐host plants that disrupted host‐plant finding by the cabbage root fly, fewest eggs (18% of check total) were laid on host plants surrounded by the weed Chenopodium album L., and most (64% of check total) on those surrounded by the weed Fumaria officinalis L. Of the 15 plants that disrupted host‐plant finding in the preliminary tests involving the onion fly, the most disruptive (8% of check total) was a green‐leaved variant of the bedding plant Pelargonium × hortorum L.H. Bail and the least disruptive (57% of check total) was the aromatic plant Mentha piperita × citrata (Ehrh.) Briq. Plant cultivars of Dahlia variabilis (Willd.) Desf. and Pelargonium×hortorum, selected for their reddish foliage, were less disruptive than comparable cultivars with green foliage. The only surrounding plants that did not disrupt oviposition by the cabbage root fly were the low‐growing scrambling plant Sallopia convolvulus L., the grey‐foliage plant Cineraria maritima L., and two plants, Lobularia maritima (L.) Desv. and Lobelia erinus L. which, from their profuse covering of small flowers, appeared to be white and blue, respectively. The leaf on which the fly landed had a considerable effect on subsequent behaviour. Flies that landed on a host plant searched the leaf surface in an excited manner, whereas those that landed on a non‐host plant remained more or less motionless. Before taking off again, the flies stayed 2–5 times as long on the leaf of a non‐host plant as on the leaf of a host plant. Host‐plant finding was affected by the size (weight, leaf area, height) of the surrounding non‐host plants. ‘Companion plants’ and aromatic plants were no more disruptive to either species of fly than the other plants tested. Disruption by all plants resulted from their green leaves, and not from their odours and/or tastes.     

Inter- and intra-population differences in the effects of temperature on postdiapause development of Delia radicum

Canadian populations of D. radicum differ in their response to temperature during postdiapause development. Populations that are primarily of the early-emerging type ( ) (St-Jean, Quebec; London, Ontario) have high values for the parameters describing this response: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaamOuamaaBa% aaleaacaWGTbaabeaakiabg2da9aaa!38F2!$$R_m = $$ 12.7–13.3; 28.0–31.8 °C ; T = 10.3–14.2 ( , the maximum developmental rate at the temperature, [ °C ] where the developmental rate is highest, and T , the parameter which gives the shape of the truncated normal curve fitted to the data), a low degree-day requirement for emergence (160–232 ), and may lack a developmental delay at temperatures above ca. 21 °C . Populations of the late-emerging type (Kildare, Prince Edward Island) have low parameter values ( , °C ; T = 6.4), high degree-day requirements (530 ), and a developmental delay at high temperatures. The parameters for the early-emergers in the population from Winnipeg, Manitoba (74% early) were intermediate ( , °C , T = 10.7, ), but resembled the early rather than the late type. This population varied from 31 to 90% early type over a 10-year period and the rate of postdiapause development at 20 °C was directly related to the percentage early. In the year with the most rapid development (90% early), development was significantly slower than in the populations from other locations with predominantly early populations, and the year with the slowest development (31% early) showed significantly faster development than that from Kildare, Prince Edward Island (100% late). Therefore the parameters for early and late types of development will not be accurate for use in mixed populations, and the parameters in mixed populations will change among years. Populations of D. radicum in North America and Europe (67 locations by years) varied from 0–100% early. At Winnipeg, the percentage early was directly related to the annual temperature accumulation ( ) during the growing season. The calculation of developmental parameters for the early-emergers of mixed populations provides a more accurate basis for estimating the times of first emergence and the first peak of emergence than parameters based on the whole population. Since postdiapause developmental rates vary both among and annually within locations, developmental models should be designed to include such variations.     

Upwind flight by the cabbage root fly, Delia radicum

ABSTRACT. In fourteen releases, most female D.radicum (L.) (Diptera, Anthomyiidae) flew upwind or at an angle to it of less than 77° regardless of the presence of host-plant odour. Females ready to lay eggs flew upwind without prior stimulation by odours from either a host crop or a trap releasing up to 3 ml/day of the attractant allylisothiocyanate. Upwind flight was more pronounced in flies from a diapause than from a continuous, non-diapause culture. Males from the non-diapause culture dispersed upwind and downwind in more or less equal numbers; old males flew mainly downwind. But, like the females, most males from the diapause culture flew upwind. Long-distance, odour-modulated anemotaxis did not appear to be used to locate distant host crops by either sex. It is concluded that the distances of insect orientation to plant odours recorded to date are only of intermediate range, and that long-range orientation to the odours of a host-plant still has to be proven. It is suggested that host-plant volatiles are involved not only in the final stage of host location but also in the first, and probably most important stage of host selection whilst the insect is still in flight.     

Infochemicals mediating the foraging behaviour of Aleochara bilineata Gyllenhal adults: sources of attractants

Aleochara bilineata Gyll. (Coleoptera: Staphylinidae) adults feed on eggs and larvae of cabbage maggot, Delia radicum (L.) (Diptera: Anthomyiidae), and might be used to control this pest. We undertook tests in a Y-tube olfactometer to determine if infochemicals are involved in the food foraging behaviour of A. bilineata adults and to precisely determine the potential sources of stimuli. A. bilineata adults oriented toward the stimuli from both the food-plant of the prey (rutabaga: Brassica napus var. napobrassica (L.) Reichb.) and the prey-plant complex (rutabaga infested by cabbage maggot larvae), but adults significantly preferred the effluvia of infested to uninfested rutabaga in a choice test. The sources of infochemicals in the infested rutabaga were the cabbage maggot larvae themselves and their frass. However, in a choice test adults preferred the effluvia of the larvae to that of the damaged rutabaga from which those larvae were obtained. It is not expected that a generalist predator uses precise herbivore-derived signals in food foraging activity, but the preference of A. bilineata adults for the larval volatiles over frass volatiles may reflect the dependence of this species on dipteran pupae for reproduction.     

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.     

Effect of expected offspring survival probability on host selection in a solitary parasitoid

Optimal Foraging Theory predicts that parasitoid females should optimize their host selection to maximize their lifetime fitness gain and parasitize the most profitable hosts. In particular, in solitary parasitoids, females should avoid superparasitism, at least when sufficient unparasitized hosts are available. However, when unparasitized hosts are scarce, they should prefer, among already parasitized hosts, those that provide the best survival probability to their progeny, which depends on the age and the developmental stage of the first parasitoid. To test this hypothesis in a solitary ectoparasitoid, Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae), we first assessed the survival probability of a second parasitoid according to the time elapsed since initial parasitism. We then analyzed the female selection behavior in patches containing a mixture of hosts parasitized over various time intervals. Our results showed that the older the opponent larva was, the lower the survival probability of the second parasitoid was. However, when the first individual had reached the prepupal stage, both individuals could complete their development. At this stage, the survival probability of the second parasitoid was surprisingly high but such individuals were reduced in size. Our study also showed that host acceptance by females was strongly correlated with the survival probability of their progeny when the first parasitoid was from 0 to less than 10 days‐old. When the first parasitoid had reached the prepupal stage, females usually rejected these hosts, although the survival probability of the offspring was quite high. This discordance between female host selection behavior and progeny survival probability is discussed.     

Distant olfactory response of the onion fly, Delia antiqua, to host-plant odour in the field

ABSTRACT. Distant olfactory orientation of female adult Delia antiqua (Meigen) to the host-plant Allium volatile dipropyl-disulphide (DPDS) was examined in the field using mark-release-recapture experiments and observations of flight behaviour. Onion-reared, post-diapause, virgin females from a laboratory colony dispersed upwind when released in the centre of 25, 50 and 100 m radius circles of eight 50 μl UDPDS baits. Percentage recapture and dispersal directedness did not decrease as a function of increasing distance to baits. In all cases the mean flight direction of recaptured flies closely correlated with mean wind direction. However, modes of the circular distributions of recaptured flies were located further crosswind when odour-baits were more distant. When distance was held constant (25 m) and DPDS concentration serially reduced (500–0.05 μ/bait), flies dispersed randomly in the absence of DPDS, crosswind in response to 0.05 μl baits and upwind in response to all other baits. Percentage recaptures on DPDS-baited traps of all concentrations were significantly greater than unbaited traps. Results from markrecapture studies were corroborated by observations of flight behaviour downwind. Flies located 100 m downwind from 50 μl DPDS baits flew upwind on take off while take-off flights in the absence of DPDS were random. Our data indicate that Allium volatiles like DPDS are involved not only in the acceptance phase of host-selection, but also in the first and probably most important stage when onion flies are initiating search long distances downwind. We conclude that D. antiqua orients to host-plants using olfactory cues from distances that should be classified as long-range ( sensu Kennedy , 1977     

Influence of background on host-plant selection and subsequent oviposition by the cabbage root fly (Delia radicum)

The behaviour of female cabbage root flies during host plant selection was studied in the laboratory using brassica plants growing in backgrounds of bare soil, clover, grass, peas and four non-living materials. Gravid females landed about twice as often on brassica plants growing in bare soil than on comparable plants growing amongst non-host plants. Once a receptive female landed on a brassica plant, the female made, on average, four ‘spiral flights’ and two jumps on and off the plant before laying alongside the plant. Surrounding a brassica plant with a diverse background altered the behaviour of the flies, so that the spiral flights around the host plant were replaced by short hops between nearby vertical objects. The loss of contact and recontact with the host plant then prevented the females from accumulating sufficient contacts with the host plant to be stimulated to lay. Spiral flights around host plants appear to determine whether or not flies will lay alongside host plants. Flies in mixed plantings have a reduced rate of settling on the host plant, and a higher rate of locomotion, because they land frequently on non-host plants. Hence, visual stimuli appeared to have greater effects than, chemical or mechanical barriers in deterring flies from laying alongside brassica plants in diverse backgrounds. In ‘choice’ situations, backgrounds of real plants reduced oviposition alongside brassica plants by at least 50%. In ‘no-choice’ situations, flies laid similar numbers of eggs alongside all brassica plants irrespective of plant background or plant size. If numbers of fly eggs are to be reduced on commercial brassica crops by undersowing the crops with clover, plants growing in bare soil may also have to be included to provide the flies with sites preferred for oviposition.     

Oviposition and chemosensory stimulation of the root flies Delia radicum and D. floralis in response to plants and leaf surface extracts from resistant and susceptible Brassica genotypes

In Brassica crops differences in susceptibility to root fly attack can be largely attributed to antixenotic resistance. Plants of four genotypes (two swedes and two kales) with widely differing resistance in field trials, were compared in laboratory choice assays for their susceptibility to oviposition by the root flies Delia radicum (L.) and D. floralis (Fallen) (Diptera, Anthomyiidae). For both species the preference among the genotypes corresponded to the susceptibility of the genotypes in the field. The preference ranking in response to surrogate leaves treated with methanolic surface extracts of the four genotypes was identical to the preference among potted plants, demonstrating that chemical factors on the leaf surface mediate host preference for oviposition in these species.For both species of fly, glucosinolates are major oviposition stimulants and for D. radicum an additional, nonglucosinolate oviposition stimulant, presently called CIF, is known. We describe a procedure for chromatographic separation of glucosinolates from CIF in leaf surface extracts. In oviposition-choice assays with D. radicum, the CIF-fractions of the two swede genotypes applied to surrogate leaves received a 1.8 and 4.6 times higher proportion of eggs than the respective glucosinolate-fractions, confirming the major importance of CIF as an oviposition stimulant. The genotype of swede that was preferred by both fly species in tests with plants and methanolic leaf surface extracts, also stimulated oviposition more in tests with the glucosinolate-fractions or the CIF-fractions derived from the surface extracts, respectively. Thus, glucosinolates and CIF together account for the observed preference among the genotypes and may also be responsible for their susceptibility under field conditions. In the two kale genotypes the preference for plants or surface extracts differed from the preference among the corresponding glucosinolate- and CIF-fractions, indicating that additional, as yet unknown chemical factors may also be involved.For both groups of stimulants tarsal chemoreceptors allow electrophysiological monitoring of glucosinolate- and CIF-activity in fractionated surface extracts. For D. radicum the chemosensory activity of both glucosinolate- and CIF-fractions corresponded to the respective behavioural activity in the oviposition preference tests, suggesting that preference for oviposition among genotypes can be predicted from the electrophysiological activity of their fractions. The chemosensory response of D. floralis, in particular to the CIF-fractions, was less pronounced than the response of D. radicum, indicating interspecific differences in the perception of the major oviposition stimulants. We discuss the potential application of electrophysiological techniques in support of other screening methods used in breeding for root fly resistance in Brassica crops.