Ground beetle (Coleoptera: Carabidae) diversity, activity density, and community structure in a diversified agroecosystem

Diversity and abundance of ground beetles (Coleoptera: Carabidae) can be enhanced in vegetable and field intercropping systems, but the complexity of polycultures precludes the application of generalized assumptions of effects for novel intercropping combinations. In a field experiment conducted at Lacombe and Ellerslie, Alberta, Canada, in 2005 and 2006, we investigated the effects of intercropping canola (Brassica napus L.) with wheat (Triticum aestivum L.) on the diversity and community structure of carabid beetles, and on the activity density responses of individual carabid species. Shannon-Wiener diversity index scores and species evenness increased significantly as the proportion of wheat comprising total crop plant populations increased in one site-year of the study, indicating a positive response to enhanced crop plant species evenness in the intercrops, and in that same site-year, ground beetle communities in intercrops shifted to more closely approximate those in wheat monocultures as the percentage of wheat in the intercrops increased. Individual carabid species activity densities showed differing responses to intercropping, although activity densities of some potential root maggot (Delia spp.) (Diptera: Anthomyiidae) predators were greater in intercrops with high proportions of wheat than in canola monocultures. The activity density of Pterostichus melanarius (Illiger), the most abundant species collected, tended to be greater in canola monocultures than high-wheat intercrops or wheat monocultures. We conclude that intercrops of canola and wheat have the potential to enhance populations of some carabid species, therefore possibly exerting increased pressure on some canola insect pests.     

Insect-crop interactions in a diversified cropping system: parasitism by Aleochara bilineata and Trybliographa rapae of the cabbage root fly, Delia radicum, on cabbage in the presence of white clover

Parasitism of the cabbage root fly, Delia radicum (L.) by the staphylinid Aleochara bilineata Gyllenhal and the cynipid Trybliographa rapae Westwood was examined in a cabbage monoculture and a mixed stand of cabbage undersown with white clover. Number of overwintering cabbage root fly pupae per plant was consistently reduced in the mixed stand, and the incidence of plants attacked by cabbage root fly was either reduced or not different in the mixed stand compared to cabbage monoculture. For both parasitoids, the probability of D. radicum attacked plants having at least one parasitized pupa increased with density of cabbage root fly pupae around the plant. For A. bilineata, this positive relation between presence of parasitism and host density was consistently stronger in cabbage monoculture than in cabbage undersown with clover. Location of a host plant by T. rapae was not consistently affected by the presence of clover. D. radicum attacked plants situated in the cabbage and clover mixture were found by T. rapae as easily as in cabbage monoculture. Overall, the total risk of parasitism for a cabbage root fly pupa by A. bilineata was reduced in the mixed stand compared to the cabbage monoculture, whereas the risk of parasitism by T. rapae was not consistently affected by clover. For both parasitoids, intensity of parasitism showed a variable relationship with host density on individual plants attacked by the cabbage root fly. Overall, in spite of consistently lower total density of pupae in the mixed cabbage—clover than in cabbage monoculture, the density of unparasitized pupae was reduced by the presence of non-host plants only in two of the four experiments. The results emphasize the need to include not only herbivore and crop, but also other plant species as well as natural enemies when evaluating management methods.     

Responses of the specialist biological control agent,Aleochara bilineata,to vegetational diversity in canola agroecosystems

Plant biodiversity is known to affect insect populations, both herbivores and their natural enemies, and as a consequence, habitat management through increased plant species composition and abundance can be exploited for sustainable pest management. In agroecosystems where crop monocultures are the routine production practice, plant biodiversity can be increased by maintaining small populations of weeds, with potential beneficial effects arising from concomitant increases in the abundance of predator and parasitoid populations. We manipulated weed populations in both species of canola, Brassica rapa L. and Brassica napus L., to investigate responses of adults of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), an important natural enemy of root maggots (Delia spp., Diptera: Anthomyiidae). Larval root maggots feed on canola taproots, disrupting the flow of water and nutrients within the plants, causing substantial yield reductions. Aleochara bilineata is a predator–parasitoid that attacks all root maggot pre-imaginal life stages. Activity density of A. bilineata increased as monocotyledonous weed biomass declined. Significant preferences between canola species were observed, with A. bilineata associated most frequently with B. rapa compared with B. napus. Our research suggests that improved management of root maggot infestations in canola through enhancement of populations of the A. bilineata predator–parasitoid could be accomplished by reducing weed infestations; however, such recommendations should consider other predators in the system and the role of weeds in reducing root maggot oviposition and damage.     

Influence of a bimodal emergence strategy of a Dipteran host on life-history traits of its main parasitoids

Abstract. 1. Among a great diversity of other strategies, insects have evolved polymodal emergence patterns that can increase survival in the face of annual variations in environmental conditions. Delia radicum L. (Diptera: Anthomyiidae) is a polyvoltine pest that attacks several cultivated cruciferous species. At the time of emergence, most populations show a polymodal emergence of type A (i.e. all individuals that enter diapause in the same year terminate diapause the following growing season, and exhibit a bimodal emergence curve). This results in the occurrence of two sympatric phenotypes, early and late, which differ by the timing of adult emergence in both diapausing and non‐diapausing generations. 2. In Brittany, D. radicum pupae can be heavily parasitised by three parasitoids, Aleochara bilineata Gyll. (Coleoptera: Staphylinidae), Aleochara bipustulata L., and Trybliographa rapae (Hymenoptera: Eucoilidae). As in all parasitoids species, the successful development of both Aleochara species and T. rapae larvae depends mainly on the amount and quality of the food provided by the host. 3. The relationship between early and late host phenotype and (i) the parasitism efficiency, (ii) emergence patterns, and (iii) host selection behaviour were investigated for the three parasitoid species. 4. Depending on parasitoid species and their different parasitism and development mode, the results reveal that host phenotype can influence (i) survival and development time in T. rapae, and (ii) survival in Aleochara species. Aleochara larvae did not appear to discriminate between early and late host pupae on the basis of phenotype, but rather selected them according to their developmental stage. Furthermore, it was discovered that the same phenological strategy occurred in T. rapae and in D. radicum. However, for D. radicum the results indicate that such a strategy has a cost as the longer development time of late host pupae results in a longer period of time favourable for parasitism.     

Life history comparison between two competitive Aleochara species in the cabbage root fly, Delia radicum: implications for their use in biological control

This study evaluates the efficacy of Aleochara bilineata Gyll and Aleochara bipustulata L. (Coleoptera: Staphylinidae) as biological control agents against the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). Biological and demographic characters are documented and comparisons made between the two species. The following results were obtained:(1) The reproductive potential and longevity of A. bipustulata females are significantly higher than those of A. bilineata. These differences are correlated with the biology and ecology of the hosts of both species.(2) Under laboratory conditions the rate of increase (r_m) of both staphylinids species is significantly higher than that of the pest and their major competitor Trybliographa rapae West. (Hymenoptera:Figitidae). This may indicate that A. bilineata and A. bipustulata can be effective biological control agents.(3) Aleochara bilineata appears to be the most interesting biological control agent against D. radicum, because of its higher rate of increase, higher host specificity and host acceptance, and a development time which is well synchronised with that of its host. With its higher competitive ability, a population of A. bilineata may increase even if competition occurs. However, A. bipustulata presents interesting biocontrol characteristics, but the best strategy may be only to use the potential of its natural populations.     

Spatial patterns of Trybliographa rapae parasitism of Delia radicum larvae in oilseed rape and cauliflower

Trybliographa rapae (Westwood) is an important parasitoid of Delia radicum (L.). Parasitism of D. radicum larvae by T. rapae in relation to host density on canola (oilseed rape) and cauliflower roots was examined at 10 field sites in Germany and Switzerland. For roots with host larvae, the proportion of roots with one or more parasitized hosts increased with increasing host density. However, for these infested roots, the parasitism of individual larvae was not consistently related to host density. When considering only roots on which there were parasitized larvae and the opportunity for multiple attacks, the proportion of larvae that were parasitized decreased with increasing host density in the field locations, and in a cage study under controlled conditions. A model of patch‐finding and number of attacks by female parasitoids suggests that patch‐finding is density‐dependent, but that low attack rate and interference effects limit numbers of attacks to three or less per visit to a host patch; the reduced number of attacks per visit leads to the inverse relationship of larval parasitism with host density in the host patches visited. The interplay of the density‐dependent and inversely density‐dependent processes appears to be responsible for the inconsistency of density dependence of overall larval parasitism in this and previous studies. In the laboratory, adult female T. rapae parasitized hosts at ≤4 cm deep in soil, but not at 6 cm deep. From the depth distribution of larval feeding sites in the field, we infer that between 4% and 20% of Delia larvae may be in a physical refuge from T. rapae parasitism, which may have a stabilizing influence on the host–parasitoid interaction.     

Aleochara suffusa andA. bilineata (Col.: Staphylinidae) as parasitoids of brassica root flies in northern Norway

A sample ofDelia puparia collected in late autumn from a brassica field at Tromsø, northern Norway, was investigated to study the level of parasitism byAleochara. BothA. suffusa andA. bilineata were reared from puparia of the cabbage root fly,Delia radicum, and the bean seed flies,D. florilega and/orD. platura. Only two specimens ofA. bilineata emerged from puparia of the turnip root fly,D. floralis. BothAleochara species hibernated in the larval state and both pupated inside the host puparium. Most specimens ofA. suffusa emerged from small hosts (D. florilega/D. platura), whereas the majority of A.bilineata emerged from host species of larger size (D. radicum/D. floralis). The time to develop from first instar larva to adult was similar for bothA. suffusa andA. bilineata. Parasitoids developing in large hosts emerged later than those in small hosts, the delay being the same for both species ofAleochara.     

Interactions between an eucoilid [Hymenoptera] and a staphylinid [Coleoptera] parasitoid of the cabbage root fly

The interaction betweenTrybliographa rapae andAleochara bilineata, 2 parasitoids of the cabbage root fly, is discussed. Larvae ofA. bilineata could not differentiate between cabbage root fly pupae containingT. rapae in its endoparasitic state and unparasitized pupae but could recognize pupae containingT. rapae once the latter had reached its ectoparasitic state. Attack byA. bilineata whileT. rapae was still in its endoparasitic state usually resulted in the staphylinid killing the eucoilid. IfT. rapae had reached the ectoparasitic state before the host pupa was attacked byA. bilineata larvae the eucoilid survived attack by the beetle larva. Multiparasitism, however, resulted in increased levels of mortality of both parasitoid populations.      

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.     

Behavioural responses to dimethyl disulphide by Aleochara bilineata and Aleochara bipustulata

Adult Aleochara bipustulata L. and Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae) are predatory on immature stages of cabbage root fly Delia radicum (L.) (Diptera: Anthomyiidae). Larvae of the two Aleochara are parasitoids of D. radicum pupae. Female Aleochara lay eggs near D. radicum puparia; the newly‐hatched Aleochara larvae enter puparia and consume the contents. Delia radicum‐infested roots of brassicas give off dimethyl disulphide (DMDS). In the field, DMDS attracts adult Aleochara to pitfall traps but does not enhance the biological control of D. radicum. In the present study, we investigate the behavioural responses of the Aleochara to DMDS in still air, as well as in moving air in a Y‐tube olfactometer, and also investigate the influence of DMDS on host selection. In larvae of both Aleochara species, DMDS induces a restricted‐area search in still air, resulting in elevated frequencies of attack of D. radicum puparia close to a source of DMDS. In the olfactometer, newly‐emerged virgin adults of both sexes of both Aleochara species choose alternatives to DMDS, older recently‐mated females are attracted to DMDS, and older males and older mate‐deprived females show no preference. Mating status of males determines the switch of their response to DMDS from avoidance to indifference. We conclude that DMDS is an important cue for host‐finding, although other cues are involved in mate‐finding. We discuss the implications for use of DMDS to enhance D. radicum mortality and for parasitism of nontarget species if A. bipustulata is introduced to Canada for biological control of D. radicum.     

Effects of conservation strip and crop type on natural enemies of Delia radicum

This 3‐year field study investigated the combined effect of floral resources and perennial shelter habitats (i.e. conservation strips), and crop rotation in supporting natural enemies of the cabbages root fly, Delia radicum. Habitat manipulation with conservation strips increased the overall catches of hymenopteran parasitoids. However, conservation strips did not increase parasitism by either of the two dominant parasitoid species, Trybliographa rapae and Aleochara bipustulata, in any study year. In fact, higher parasitism was found in control plots in the second year. This could be explained by parasitoid mobility and higher patch detectability, as more plants in the control plots were infested with D. radicum larvae. Conservation strips did not result in increased predation of D. radicum eggs. However, the activity densities of two Bembidion species were correlated with egg predation. The species assemblage distribution of epigeal predators was best explained by seasonal period, followed by year and, to a low extent, crop type, while treatment with conservation strips had no effect. However, during the egg laying peak of D. radicum, a higher number of A. bipustulata, an important larval predator was observed in conservation strips during one study year. In conclusion, positive effects of conservation strips were demonstrated for abundance of some natural enemies of D. radicum, but a consistent increase in performance could not be adequately demonstrated due to experimental set‐up, the short timescale and the complex landscape in which our study site was located.     

Host range testing of a prospective classical biological control agent against cabbage maggot,Delia radicum,in Canada

The introduction of a European natural enemy, Aleochara bipustulata L. (Coleoptera: Staphylinidae), is being considered for control of cabbage maggot, Delia radicum (L.) (Diptera: Anthomyiidae) in canola in Canada, and the host specificity of this pupal parasitoid must first be studied. Contemporary guidelines were used to select 18 species of Diptera to represent non-target species taxonomically related or ecologically similar to reported hosts of A. bipustulata, or beneficial species. No-choice tests were used to determine which of the 18 species are within A. bipustulata’s fundamental host range, and whether puparial structure or mass or duration of pupal development influence their acceptability and suitability as hosts. Five species were consistently suitable as hosts, and these were either relatively small or were taxonomically closely related to the target host. The probability that a puparium would be entered by a parasitoid larva was greatest for small puparia, but was unaffected by pupal duration. The probability of completing parasitoid development once a puparium was entered was influenced by both puparial mass and duration of pupal development. Pitfall traps to assess habitat associations caught A. bipustulata adults in a variety of crop habitats but none were caught in forests. Host range and habitat data are used to argue that there is little risk of parasitism to beneficial taxa. Non-target species taxonomically closely related to the target D. radicum or with small puparia may fall within the fundamental host range of A. bipustulata. However, risk to many of these species may be minimal because of the habitat preferences of the parasitoid and its cues for host-finding and recognition.     

Toxicity of the insect growth regulator diflubenzuron to the rove beetle Aleochara bilineata, a parasitoid and predator of the cabbage maggot Delia radicum

The insect growth regulator Dimilin^®, common name: diflubenzuron, applied at concentrations effective against the cabbage maggot Delia radicum L., did not affect the hatching of eggs of the staphylinid predator and parasite, Aleochara bilineata Gyllenhal. Applied externally in distilled water, Dimilin was non-toxic to first instar larvae, to larval instars developing within host puparia, and to adults. When Dimilin was dissolved in dimethylsulfoxide, then applied to the exoskeleton of host puparia, emergence of A. bilineata adults was suppressed. The conventional insecticide Furadan, at a concentration (0.001% in distilled water) equal to the lowest concentration of Dimilin used, was toxic to both eggs and larvae of A. bilineata.

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Résumé Lorsqu' appliqué dans des concentrations efficaces contre la mouche du chou Delia radicum L. (0.1, 0.01, 0.001% p/v dans de l'eau distillée), le Dimilin (diflubenzuron), un régulateur de croissance des insectes, n'a pas affecté l'éclosion des oeufs de Aleochara bilineata Gyllenhal, un staphylin prédateur et parasite. En application externe, à la plus forte concentration (0.1% p/v dans de l'eau distillée), cet inhibiteur de synthèse de la chitine s'est avéré non toxique pour les larves du premier stade, les stades larvaires qui se développent à l'intérieur du puparium de l'hôte, et les adultes. Le traitement au Dimilin n'a pas affecté la fertilité des staphylins adultes. Cependant, lorsque le diflubenzuron était dissous dans du diméthylsulfoxyde et appliqué sur l'exosquelette du puparium de l'hôte, l'émergence des adultes de A. bilineata était éliminée. L'insecticide conventionnel Furadan (carbofurane), à une concentration égale à la plus faible concentration de Dimilin utilisée (0.001% p/v dans de l'eau distillée), s'est avéré toxique à la fois pour les oeufs et pour les larves de A. bilineata.

     

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.     

Effects of rhizobial inoculation, cropping systems and growth stages on endophytic bacterial community of soybean roots

Interspecific interactions and soil nitrogen supply levels affect intercropping productivity. We hypothesized that interspecific competition can be alleviated by increasing N application rate and yield advantage can be obtained in competitive systems. A field experiment was conducted in Wuwei, Gansu province in 2007 and 2008 to study intercropping of faba bean/maize, wheat/maize, barley/maize and the corresponding monocultures of faba bean (Vicia faba L.), wheat (Triticum aestivum L.), barley (Hordeum vulgare L.) and maize (Zea mays L.) with N application rates of 0, 75, 150, 225 and 300 kg N ha^?1. Total land equivalent ratios (TLER) were 1.22 for faba bean/maize, 1.16 for wheat/maize, and 1.13 for barley/maize intercropping over the 2-year study period. Maize was overyielding when intercropped with faba bean, but underyielding when intercropped with wheat or barley according to partial land equivalent ratios (PLER) based on grain yields of individual crops in intercropping and sole cropping. There was an interspecific facilitation between intercropped faba bean and maize, and interspecific competition between maize and either wheat or barley. The underyielding of maize was higher when intercropped with barley than with wheat. Fertilizer N alleviated competitive interactions in intercrops with adequate fertilizer N at 225 kg ha^?1. Yield advantage of intercropping can be acquired with adequate nitrogen supply, even in an intensive competitive system such as barley/maize intercropping. This is important when using intercropping to develop intensive farming systems with high inputs and high outputs.     

Parasitism capacity and searching efficiency of Diaeretiella rapae parasitizing Brevicoryne brassicae on susceptible and resistant canola cultivars

Different cultivars of aplant species can affect the foraging and efficiency of natural enemies, both directly through physical and biochemical properties or indirectly through the herbivore's diet. In this study, the parasitism capacity and functional response of Diaeretiella rapae McIntosh were determined on the cabbage aphid, Brevicoryne brassicae (L.) reared on susceptible (Opera) and resistant (Okapi) canola cultivars under laboratory conditions at 25?±?1?°C, 60?±?5% RH and a16:8?h L:D photoperiod. The parasitoid exhibited Type II and Type III functional responses on the resistant and susceptible cultivars, respectively. The estimated value of searching efficiency (a) was 0.1637?±?0.1095?h^?1 on the resistant cultivar whereas its value was dependent on host density on the susceptible cultivar. The handling times (T_h) on the susceptible and resistant canola cultivars were 0.108?±?0.040 and 0.320?±?0.048?h, respectively. The net parasitism rate (C₀) of the parasitoid wasp varied from 128.09 hosts per parasitoid lifetime on the susceptible to 71.01hosts on the resistant canola cultivar. The transformation rate from host population to parasitoid offspring (Qp) was equal to 1 on both cultivars (C₀?=?R₀). The finite parasitism rate (ω) on the susceptible cultivar (0.819 hosts per parasitoid per day) was significantly higher than that on the resistant one (0.578 hosts per parasitoid per day). In conclusion, canola cultivars affected the performance of D. rapae in controlled small-scale laboratory experiments and compared with the susceptible cultivar, the resistant one had anadverseeffect on the efficiency of the parasitoid.     

Comparative study of the antennal sensilla of five species of root maggots: Delia radicum L., D. floralis F., D. antiqua Mg., D. platura MG. (Diptera : Anthomyiidae) and Psila rosae F. (Diptera : Psilidae)

A comparative study of the antennal sensilla of Delia radicum L., D. floralis F., D. antiqua Mg., D. platura Mg. (Diptera : Anthomyiidae) and Psila rosae F. Diptera Psilidae) is undertaken. For both sexes of each species, the type, distribution, and density of sensilla are determined. All 5 species have trichoid (olfactory) and grooved (olfactory) sensilla. Basiconica I (blunt) sensilla (olfactory) are found on each of the species examined, except D. platura. Basiconica II (tapered) (olfactory) and clavate (olfactory) sensilla are found only on Delia species. Also, only Delia species have single-chambered, dorsal pits, and these contain basiconic II pit sensilla (olfactory). Common to all 5 species is a multi-chambered ventral pit (olfactory). In the ventral pit, all 5 species have grooved pit sensilla (olfactory). In addition to this type of sensillum the Delia species have smooth-walled conical pit sensilla (hygro-/thermosensitive) and P. rosae has granular pit sensilla (hygro-/thermosensitive). Smooth-walled tapered pit sensilla (hygro-/thermosensitive) are found in D. radicum. Similarities and differences in the density of surface sensilla between dorsal and ventral funicular surfaces, male and female flies, and oligophagous (D. antiqua, D. radicum, D. floralis and P. rosae) and polyphagous (D. platura) species are compared. Several differences in sensillum density between the dorsal and ventral funicular surfaces are observed, but these do not fit into a consistent trend. Except for D. radicum, there are differences in sensillum density between male and female flies. For the oligophagous species, females have a greater sensillum density, whilst for the polyphagous D. platura males have a greater sensillum density. Comparisons between species show the greatest differences between the Delia species and P. rosae, and within the 4 Delia species, differences in sensillum density do not correlate with host range or body size.     

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.     

Nitrogen economy in relay intercropping systems of wheat and cotton

Relay intercropping of wheat and cotton is practiced on a large scale in China. Winter wheat is thereby grown as a food crop from November to June and cotton as a cash crop from April to October. The crops overlap in time, growing as an intercrop, from April till June. High levels of nitrogen are applied. In this study, we analyzed the N-economy of the monocultures of cotton and wheat, and of four relay intercropping systems, differing in number of rows per strip of cotton or wheat. Field experiments were carried out from 2001/02 to 2003/04 in the Yellow River region in China. We quantified the nitrogen uptake and nitrogen use efficiency of wheat and cotton in relay intercropping systems to test if intercrops are more resource use efficient in comparison to monocrops. Nitrogen (N) yields of wheat per unit area in the four intercropping systems were lower than in the monocrop, which ranged from 203 to 288 kg ha⁻¹. The total N-uptake per unit biomass was similar between wheat in mono- and intercrops. On average, the N-yield of cotton per unit area was lower in intercrops than in monocrops, which ranged from 110 to 127 kg ha⁻¹, but the total N-uptake per unit biomass was higher in intercropped cotton, as dry matter production was reduced to a greater extent by intercropping than N-uptake. The N-uptake of cotton was diminished during the intercropping phase, but recovered partially during later growth stages. The physiological nitrogen use efficiency (IE) of wheat was not much affected by intercropping, but it was reduced in cotton, due to delayed flowering and less reproductive growth. Total N-efficiency of the system was assessed by comparing the relative nitrogen yield total (RNT), i.e. the sum of the ratio’s of total N-uptake by a component crop in the intercrop relative to the N-uptake in the monocrop, to the relative yield total. RNT ranged from 1.4 to 1.7, while the relative yield total (RYT) ranged from 1.3 to 1.4, indicating that intercrops used more nitrogen per unit production than monocrops. An analysis of the crop nitrogen balance showed that the nitrogen surplus of sole crops amounted to 220 kg ha⁻¹ for wheat and 140 kg ha⁻¹ for cotton, while in the intercropping systems, the annual N surplus exceeded 400 kg ha⁻¹. Conventional N-management in intercrops thus results in high N-surpluses that pose an environmental risk. The N management could be improved by means of a demand-based rate and timing of N applications.     

Root herbivory reduces growth and survival of the shoot feeding specialist Pieris rapae on Brassica nigra

Plants may respond to herbivore attacks by changing their chemical profile. Such induced responses occur both locally and systemically throughout the plant. In this paper we studied how Brassica nigra (L.) Koch (Brassicaceae) plants respond to two different root feeders, the endoparasitic nematode Pratylenchus penetrans Cobb (Tylenchida: Pratylenchidae) and the larvae of the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae). We tested whether the activities of the root feeders affected the survival and development of the shoot feeding crucifer specialist Pieris rapae (L.) (Lepidoptera: Pieridae) via systemically induced changes in the shoots. Overall, P. rapae larvae grew slower and produced fewer pupae on plants that were infested with root feeders, especially on plants infested with P. penetrans. This effect could not be attributed to lower water or protein levels in these plants, as the percentage of water in the controls and root infested shoots was similar, and protein content was even higher in root infested plants. Both glucosinolate as well as phenolic levels were affected by root feeding. Initially, glucosinolate levels were the lowest in root infested plants, but on P. penetrans infested plants they increased more rapidly after P. rapae started feeding than in controls or D. radicum infested plants. Plants with D. radicum feeding on their roots had the highest phenolic levels at all harvest dates. Our results indicate that root feeding can significantly alter the nutritional quality of shoots by changes in secondary metabolite levels and hence the performance of a specialist shoot feeder.