Behavioural responses of the aphid parasitoid Diaeretiella rapae to volatiles from Arabidopsis thaliana induced by Myzus persicae

In response to herbivory by insects, several plant species have been shown to produce volatiles that attract the natural enemies of those herbivores. Using a Y‐tube olfactometer, we investigated responses of the aphid parasitoid Diaeretiella rapae MacIntosh (Hymenoptera: Aphidiidae) to volatiles from Arabidopsis thaliana Columbia (Brassicaceae) plants that were either undamaged, infested by the peach‐potato aphid, Myzus persicae Sulzer (Homoptera: Aphididae), or mechanically damaged, as well as to volatiles from just the aphid or its honeydew. In dual‐choice experiments, female D. rapae given oviposition experience on A. thaliana infested with M. persicae were significantly attracted to volatiles from A. thaliana infested with M. persicae over volatiles from undamaged A. thaliana and similarly were significantly attracted to plants that had been previously infested by M. persicae, but from which the aphids were removed, over undamaged plants. Diaeretiella rapae did not respond to volatiles from M. persicae alone, their honeydew, or plants mechanically damaged with either a pin or scissors. We conclude that an interaction between the plant and the aphid induces A. thaliana to produce volatiles, which D. rapae can learn and respond to. Poor responses of D. rapae to volatiles from an A. thaliana plant that had two leaves infested with M. persicae, with the two infested leaves being removed before testing, suggested the possibility that, at this stage of infestation, the majority of volatile production induced by M. persicae may be localized to the infested tissues of the plant. We conclude that this tritrophic interaction is a suitable model system for future investigations of the biochemical pathways involved in the production of aphid‐induced volatiles attractive to natural enemies.     

Ecology of the aphid pests of protected pepper crops and their parasitoids

Myzus persicae, Macrosiphum euphorbiae, Aphis gossypii and Aulacorthum solani (Homoptera: Aphididae) are principal pests of protected pepper crops in southeastern Spain. Our goal was to determine the incidence of aphids on pepper crops and the role of vegetation surrounding greenhouses as a source of aphids and their parasitoids. The population dynamics were followed in six commercial greenhouses during 3 years. Another 82 greenhouses and their surrounding vegetation were surveyed occasionally. Myzus persicae had the highest incidence in pepper greenhouses followed by M. euphorbiae and A. solani. Parasitism of all aphid species in greenhouses was low, Aphidius matricariae and Aphidius colemani being the most abundant parasitoids. Myzus persicae and Macrosiphum euphorbiae were the most abundant and polyphagous aphids, being present on 77 and 55% of the plants sampled outside greenhouses, respectively; species of Brassicaceae were the main hosts for both aphids. Aulacorthum solani was only present on Malva parviflora and at low numbers. Outside greenhouses, A. matricariae was the most common parasitoid of M. persicae, followed by Diaeretiella rapae and A. colemani. Aphidius matricariae was the most polyphagous, being present in 10 out of 22 aphid species. Macrosiphum euphorbiae and A. solani were both parasitised by A. ervi and Praon volucre. Aphelinus asychis was found on A. solani. Parasitoids were found in other aphids not attacking pepper. The role of natural vegetation as a reservoir of aphid pests of pepper and of parasitoids is discussed.     

Infectivity of entomopathogenic nematode Steinernema carpocapsae Pocheon strain (Nematoda: Steinernematidae) on the green peach aphid Myzus persicae (Hemiptera: Aphididae) and its parasitoids

Infectivity of entomopathogenic nematode (EPN) Steinernema carpocapsae Pocheon strain on the green peach aphid Myzus persicae and its parasitic wasps (e.g., Aphidius colemani, Aphidius gifuensis and Diaeretiella rapae) was evaluated under laboratory conditions. Infective juveniles (IJs) of S. carpocapsae Pocheon strain had low infectivity against nymph and adult stages of M. persicae, showing 2% and 6.7% of mortality, respectively. Application of the EPNs had little effect on mummies caused by the three parasitoid species, allowing them to remain intact. No IJ invaded the host, regardless of EPN application rate. The parasitoid emergence from mummies ranged from 80% to 85% in the presence of EPN while 79–86% was recorded in the absence of EPN. However, the presence of the IJs reduced oviposition by the three parasitoid species, decreasing the rate up to 59% when the nematodes were applied before parasitoid release, while little difference in oviposition was observed when nematodes were applied after parasitoid release.     

Low parasitism by Diaeretiella rapae (Hym.: Braconidae) of Lipaphis pseudobrassicae (Hemip.: Aphididae): pre- or post-ovipositional host resistance?

While the aphid Lipaphis pseudobrassicae (Davis) (Hemip.: Aphididae: Macrosiphini) is considered one of the preferred hosts of Diaeretiella rapae (McIntosh) (Hym.: Braconidae: Aphidiinae) in several parts of the world, field surveys in Uberlandia (Brazil) found parasitism of this aphid to not exceed 10%. This study sought to determine the cause of this low parasitism, as well as the effects of parasitism on the intrinsic growth rate of the aphid population. We evaluated parasitism, percentage emergence, developmental time, longevity, number of attacks and number of parasitoid larvae in L. pseudobrassicae and compared these to the same characteristics in Brevicoryne brassicae (L.) and Myzus persicae (Sulzer). The lowest percentage of parasitism was found in L. pseudobrassicae, followed by M. persicae and B. brassicae. The ratio between the number of parasitoid larvae and the number of ovipositions in L. pseudobrassicae ranged from 0.02 to 0.03, while, in B. brassicae, it was between 0.41 and 0.44 and, in M. persicae, between 0.62 and 0.80, indicating high mortality rates of early stages of D. rapae in L. pseudobrassicae. Parasitism by D. rapae reduced the r_m of L. pseudobrassicae. The r_m for parasitised aphids was only 63% of that for unparasitised aphids. However, no hosts died before reaching adulthood, and 83% of parasitised aphids were still able to reproduce. As a result, the r_m of the aphid was positive, resulting in population growth of L. pseudobrassicae, even among individuals parasitised during the second instar. Our results indicate the existence of L. pseudobrassicae genotypes that are completely resistant to D. rapae.     

Host-range study about four aphid parasitoid species among 16 aphid species for constructing banker–plant systems

To provide fundamental information for the biological control of aphids in vegetable greenhouses, we compared the host ranges of four aphid parasitoid species, Aphidius colemani Viereck, Aphidius gifuensis Ashmead, Diaeretiella rapae (M’Intosh), and Ephedrus nacheri Quilis (Hymenoptera: Braconidae: Aphidiinae). The acceptability as host of 11 vegetable-pest aphids, Acyrthosiphon pisum (Harris), Aphis craccivora Koch, Aphis gossypii Glover, Aulacorthum solani (Kaltenbach), Brevicoryne brassicae (Linnaeus), Chaetosiphon fragaefolii (Cockerell), Lipaphis erysimi (Kaltenbach), Macrosiphoniella sanborni (Gillette), Macrosiphum euphorbiae (Thomas), Myzus persicae (Sulzer), and Uroleucon formosanum (Takahashi), in addition to five aphid species, Melanaphis sacchari (Zehntner), Rhopalosiphum maidis (Fitch), Rhopalosiphum padi (Linnaeus), Schizaphis graminum (Rondani), and Sitobion akebiae (Shinji) (Hemiptera: Aphididae) that serve as alternative hosts in banker–plant systems for the four aphid parasitoid species, were investigated. A newly emerged pair of parasitoid adults were provided to 100 aphids of each species on caged host plants in a 25 °C chamber for 24 h. The numbers of mummified aphids and emerged adults were counted in 10 trials for each aphid species. Aphidius colemani, A. gifuensis, D. rapae and E. nacheri parasitized four, two, three, and eight pest species, respectively, and four, three, three, and five alternative host species, respectively. Ephedrus nacheri had the broadest host range among the four species, and all the four species parasitized M. persicae, R. maidis, and S. graminum. This information will be useful for selecting candidate of biological control agents for aphids and for constructing banker–plant systems.

     

Nectar feeding increases exploratory behaviour in the aphid parasitoid Diaeretiella rapae (McIntosh)

Feeding on floral nectar has multiple positive effects on parasitic wasps, including increased longevity and fecundity, and in addition, nectar feeding can also alter parasitoid behaviour. To advance understanding of the effects of nectar feeding on Diaeretiella rapae (McIntosh) [Hymenoptera: Braconidae], the activities of 1‐day‐old female D. rapae with or without a prior buckwheat (Fagopyrum esculentum) nectar meal were quantified. Nectar increased searching time of D. rapae by a factor of 40 compared with individuals provided with water only and reduced the time spent stationary. The number of attacks to aphids by nectar‐fed parasitoids was not significantly (P = 0.06) higher than that of unfed D. rapae, suggesting that the conditions of the experiment facilitated host finding by ‘quiet’ parasitoids. Nevertheless, nectar feeding modified the behaviour of D. rapae in a way that parasitoids were more explorative and less inactive. This represents one additional mechanism through which nectar feeding impacts parasitoid biology and suggests that a synergy between increased host searching, increased longevity and increased fecundity should lead to an enhancement of biocontrol when D. rapae females have access to nectar in the field.     

Behavioural differences during host selection between alate virginoparae of generalist and tobacco-specialist Myzus persicae

Host plant selection and acceptance by aphids involves four consecutive steps: (1) prealighting behaviour, (2) leaf surface exploration and probing of subepidermal tissues, (3) deep probing of plant tissues, and (4) evaluation of the phloem sap. Host specialisation in aphids may involve not only different performances on potential hosts, but also different strategies for host selection and acceptance. Myzus persicae s.s. (Sulzer) (Homoptera: Aphididae) is one of the most polyphagous aphid species, although a tobacco‐adapted subspecies, M. persicae nicotianae, has been described. These two taxa constitute a good system for studying the effect of host range on host selection strategies. We studied the first two steps in the host selection process by alate virginoparae of M. persicae s.s. and M. persicae nicotianae on host and non‐host plants, using three types of behavioural assays: wind tunnel, olfactometry, and video‐recording. Alate virginoparae of M. persicae nicotianae recognised and chose their host plant more efficiently than M. persicae s.s., on the basis of olfactory and visual cues, and factors residing at cuticular and subcuticular levels. Host recognition was evident before phloem tissues were contacted. Olfactory cues were apparently not involved in host selection by M. persicae s.s.     

Efficacy of the parasitoid,Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae) against Myzus persicae (Sulzer) (Hemiptera: Aphididae) infesting rapeseed-mustard

Efficacy of Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae) was determined by studying its biological and behavioural attributes on the aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae). Total developmental duration of the parasitoid was longer in younger instars compared to that in older instars of aphid. Host age negatively affected the oviposition period and fecundity of the ovipositing parasitoid. Survival of the immatures was higher in older than the younger instars of the aphid. Sex ratio for the parasitoid was female-biased. Longevity of female parasitoid was higher while parasitizing aphid nymphs of younger age groups (1–2 days old) compared to that of older age groups (4–5 days old). Parasitoid showed a higher preference towards second instar of M. persicae both in choice and no-choice tests. Results on mutual interference revealed that per capita searching efficiency decreased from 0.91 to 0.07 as parasitoid densities increased from 1 to 8. Logistic regression exhibited a type II functional response for D. rapae. The estimated values of search-efficiency (a) and handling time (T_h) were 0.038 h⁻¹ and 1.59 h, respectively. The maximum parasitization rate was 15.08. This investigation suggests that D. rapae could be an effective candidate for augmentative biological control of M. persicae which infests several plant species of economic importance.     

Parasitism rate of Myzus persicae (Sulzer) by Diaeretiella rapae (McIntosh) in the presence of an alternative,resistant host

The aphids Lipaphis pseudobrassicae (Davis) and Myzus persicae (Sulzer) (Hemiptera: Aphididae) are important Brassicaceae pests, occurring worldwide and causing significant damage to crops. Interspecific variations in the resistance to natural enemies can potentially impact the interaction among aphid populations. Here we evaluated the hypothesis of associational resistance by determining if the presence of resistant aphids (L. pseudobrassicae) reduces the rate of parasitism by Diaeretiella rapae (McIntosh) on non-resistant aphids (M. persicae). The experiment was conducted using collard green plants infested with M. persicae and L. pseudobrassicae either resistant or susceptible to D. rapae. The percentage of parasitism by D. rapae was greater on L. pseudobrassicae in the susceptible than in the resistant treatment, but parasitism rates on M. persicae did not differ between the treatments. There was no difference in average growth rate between M. persicae and susceptible L. pseudobrassicae populations, but resistant L. pseudobrassicae had greater growth rate than M. persicae. These results suggest that over a short period of time the presence of resistant L. pseudobrassicae does not affect the rate of parasitism by D. rapae on M. persicae.     

Oviposition behaviour ofEphedrus cerasicola [Hym.: Aphidiidae] parasitizing different instars of its aphid host

The parasitoidEphedrus cerasicola Starý oviposited in all 4 nymphal instars and in newly moulted adults ofMyzus persicae (Sulzer). The different host categories were offered with no choice. The duration of an oviposition increased with the age of nymphs, being about 13, 18, 21, 22, and 17 s from 1 st instars to adults, respectively. Observations of number of stabbing attacks prior to oviposition, percent of the encounters not resulting in oviposition, time from first encounter to oviposition, handling time and aphid defensive behaviour also indicated that 1 st instarM. persicae are most easily parasitized. The behaviour ofE. cerasicola in encounters with unparasitized and parasitized hosts, suggested that the parasitoid could discriminate. In encounters with parasitized 1 st to 4th instar aphids,E. cerasicola used only the antennae in 80% of the encounters that resulted in discrimination.     

Interference competition between Aphidius matricariae and Praon volucre (Hymenoptera: Braconidae) attacking two common aphid species

Interference competition is a common phenomenon that occurs among adult females of different species to gain the highest food resources at the same time. In this research, extrinsic competition between Aphidius matricariae and Praon volucre on different densities and stages of two important pests of greenhouse crops in the world, Aphis gossypii and Myzus persicae, were investigated. The results of this study showed that total percentage parasitism of second, third and fourth nymphal instars of A. gossypii and M. persicae by A. matricariae, and P. volucre were affected by extrinsic competition. A combination of A. matricariae and P. volucre on A. gossypii had a negative effect on performance of these parasitoids. Although extrinsic competition reduced the efficiency of A. matricariae and P. volucre on M. persicae, total parasitism of M. persicae by A. matricariae and P. volucre (combination of two parasitoids) increased compared to A. matricariae or P. volucre acting alone. Also the results indicated that the extrinsic competition between A. matricariae and P. volucre on both A. gossypii and M. persicae was apparently more intense when presented with 25 hosts compared to 50 and 100 hosts. The results of this research show important information to determine an appropriate combination of these two parasitoid wasps for biological control of A. gossypii and M. persicae in greenhouses.     

Host foraging for differentially adapted brassica-feeding aphids by the braconid parasitoid Diaeretiella rapae

Interactions occurring in a tritrophic system comprising plants, aphids and parasitoids are of great complexity. The generalist endoparasitoid Diaeretiella rapae (McIntosh) (Hymenoptera: Aphidiidae) displays specialist characteristics on brassica feeding aphids. Previously, we studied differential signalling to D. rapae by specialist and generalist Brassicaceae feeding aphids on turnip. We reported no differences in the attractiveness of volatile compounds from the two turnip/aphid complexes. However, we reported a significantly greater D. rapae attack rate on the specialist Lipaphis erysimi (Kaltenbach) than the generalist Myzus persicae (Sulzer). As a consequence we predicted that D. rapae would forage more efficiently and produce more offspring on L. erysimi. We present here some additional data collected in a more complex spatial/temporal environment in large experimental chambers and discuss this, drawing attention to the need for careful interpretation of mechanistic information in predicting the overall foraging process.Key words: tritrophic interactions, honeydew, Lipaphis erysimi, Myzus persicae, specialist, generalist, glucosinolatePlants are a principal component of parasitoid-host interactions.¹ Plant derived chemical cues may be exploited by parasitoids to locate successfully host habitats, hosts and to assess host quality either positively or negatively.² However, there are a multitude of other direct and indirect interactions that structure aphid-parasitoid communities.³ Plants of the family Brassicaceae have a diverse and interesting secondary chemistry, which forms an important component of their defence against herbivores. This secondary chemistry includes glucosinolates located within plant cells, which upon damage are hydrolysed by enzymes known as myrosinases.^4,5 Products of these reactions include nitriles and volatile isothiocyanates. Lipaphis erysimi (Kaltenbach), the turnip aphid, is a specialist on plants of the order Capparales, particularly the Brassicaceae, and like its fellow specialist, Brevicoryne brassicae,⁶ sequesters glucosinolates using them as defence against natural enemies,⁷ whereas Myzus persicae (Sulzer), the peach-potato aphid, does not sequester glucosinolates⁷ but excretes large quantities in its honeydew.⁸ We hypothesised that Diaeretiella rapae (McIntosh) would display different foraging behaviours toward plant-derived chemical signals from these differentially adapted aphids feeding on turnip.We have studied some of the behavioural responses of D. rapae to these two aphid species on a single cultivar of turnip (Brassica rapa var rapifera cv Tokyo Cross). Volatile compounds from turnip infested by L. erysimi or M. persicae are both attractive to D. rapae, but not differentially so. The volatile composition of both turnip/aphid complexes (TAC) includes two isothiocyanates, of which one, 3-butenyl isothiocyanate, was shown to be attractive in Y-tube bioassays.⁷ In no choice tests, attack rates, defined as the number of full contacts between the ovipositor of a parasitoid and its potential aphid hosts in a minute (counting multiple stabbings repetitively on the same aphid as one attack), were significantly greater on L. erysimi than on M. persicae, irrespective of which of these aphids was the original host.⁹ This is possibly due to glucosinolate derived kairomones that may be detected either by antennation or ovipositor probing on the potential host.¹⁰ This behaviour is primed by cues received by the parasitoid during emergence from its mummy case.⁹ Taking these mechanistic results together, we hypothesised that given an equal supply of potential hosts, D. rapae would parasitize more L. erysimi aphids than M. persicae, irrespective of the host in which they developed.We tested this hypothesis in four Perspex chambers (L 1.7 m, W 1.2 m, H 1 m) with each containing four potted turnip plants infested with 250 L. erysimi aphids and four with 250 M. persicae aphids. Plants infested with the different aphid species were placed alternately within the chamber in two equally spaced parallel rows. A total of 15 naive D. rapae parasitoids were released from three points in each chamber. A total of four one-chamber replicates were made using D. rapae reared on L. erysimi and four with D. rapae reared on M. persicae. Eleven days after the release of parasitoids the number of mummified aphids on each plant was recorded.We found that D. rapae reared on L. erysimi produced significantly more mummies on L. erysimi than M. persicae, as we hypothesised. However, D. rapae reared on M. persicae produced significantly more mummies on M. persicae than L. erysimi (Fig. 1). This was contrary to our hypothesis. In addition, there were significantly more mummies produced by parasitoids reared on L. erysimi than those reared on M. persicae.Open in a separate windowFigure 1Average number of mummified aphids recovered per species (LE = Lipaphis erysimi; MP = Myzus persicae) per cage. * indicates a significant difference by t-test in the number of mummies produced on LE and MP, (t = 2.39, p = 0.048) and (t = 2.40, p = 0.048) for D. rapae reared on M. persicae and L. erysimi respectively.In an attempt to identify possible reasons for these observations we recorded different aphid patch parameters for M. persicae and L. erysimi including patch area, aphids per patch and patch density. M. persicae formed larger less dense patches than L. erysimi (Fig. 2). As well as forming different patch structures the aphids are different in colour, M. persicae being generally yellow, while L. erysimi are a dark grey/green. Green and yellow pigments reflect distinctly different spectra of light, which can be detected by the trichromatic vision common to most hymenoptera.¹⁰ Thus colonies of the different species present different visual cues, which may provide essential foraging cues to parasitoids.^11,12 However, it seems unlikely that the parasitoids used in this experiment, naive but for the conditioning received upon emergence from their mummy case, would display different responses to visual cues. We propose that the chemistry of the system is likely to hold the key to the behaviour of D. rapae, and put forward the following explanation.Open in a separate windowFigure 2Aphid patch parameters, including patch area (cm²), number of aphids per patch, and density of aphids within the patches (Aphids/cm²) for Lipaphis erysimi (LE) and Myzus persicae (MP).D. rapae does not have a preference for the volatile compounds associated with either TAC. So, though not necessarily random, it is probable that there is an equal chance of encountering each aphid species first. It is evident that one of the main differences between the two TAC is the chemical constituents and quantity of honeydew, which has been shown to alter on-plant foraging for B. brassicae by D. rapae.^13–15 The honeydew composition of aphids is species and plant specific¹⁵ and has been shown to provide parasitoids with information on host species identity through species-specific kairomones.¹⁷ M. persicae feeding on the Brassicaceae produce a honeydew rich in glucosinolates.⁸ A likely consequence is that the cuticle of the M. persicae mummy case will retain some of this chemistry which will then be received by the parasitoid upon emergence. This stage of a parasitoid''s life has been shown to influence adult responses to volatiles^18–20 and attack rate.⁴ Subsequent location of this same substance whilst foraging could prompt parasitoids to spend an increased time residing in these patches. It has been shown in the hymenopterous whitefly parasitoid Encarsia formosa (Gahan) that honeydew of a range of species including non-hosts will have an arrestant effect, but that host honeydew results in longer searching times.¹⁵ This same effect could account for our findings, and the lower total number of mummies produced could be explained by the innately lower attack rate on M. persicae compared to L. erysimi.In conclusion we draw attention to the following points.

1. It is evident that for some parasitoid species emergence from their mummy case provides information that can manifest itself in long lived host foraging decisions. We hypothesise that aphid honeydew provides an important contact cue that may influence the residence time of D. rapae, and could for other specialist parasitoids.
2. Elucidation of parasitoid host foraging mechanisms provides important information, but increasing experimental complexity may not reveal intuitive results. This is likely to become even more important under heterogeneous field conditions, highlighting the need for corroborating the significance of laboratory studies through well designed field or field-simulation studies.

     

Influence of elevated CO2 on interspecific interactions at higher trophic levels

Abstract We report the results of a study investigating the influence of elevated CO₂ on species interactions across three trophic levels: a plant (Brassica oleracea), two aphid herbivores (the generalist Myzus persicae and the specialist Brevicoryne brassicae), and two natural enemies (the coccinellid Hippodamia convergens (ladybird) and the parasitoid wasp Diaeretiella rapae). Brassica oleracea plants reared under elevated CO₂ conditions (650 ppmv vs. 350 ppmv) were larger and had decreased water and nitrogen content. Brevicoryne brassicae reared on plants grown in elevated CO₂ were larger and accumulated more fat, while there was no change in M. persicae traits. Fecundity of individual aphids appeared to be increased when reared on plants grown in elevated CO₂. However, these differences were generally lost when aphids were reared in colonies, suggesting that such changes in plant quality will have subtle effects on aphid intraspecific interactions. Nevertheless, CO₂ treatment did influence aphid distribution on plants, with significantly fewer M. persicae found on the shoots, and B. brassicae was only found on senescing leaves, when colonies were reared on plants grown in elevated CO₂. We reared B. brassicae and M. persicae in competition on plants grown at both the CO₂ concentration treatments. We found a significantly lower ratio of M. persicae: B. brassicae on plants grown under elevated CO₂ conditions, strongly suggesting that increasing CO₂ concentrations can alter the outcome of competition among insect herbivores. This was also reflected in the distribution of the aphids on the plants. While the CO₂ treatment did not influence where B. brassicae were found, fewer M. persicae were present on senescing leaves under elevated CO₂ conditions. Changes in plant quality resulting from the CO₂ treatments did not appear to alter aphid quality as prey species, as the number consumed by the ladybird H. convergens, and the number parasitised by the parasitoid wasp D. rapae, did not change. To our knowledge, this study provides the first empirical evidence that changes in host plant quality mediated by increasing levels of CO₂ can alter the outcome of interspecific competition among insect herbivores.     

Dynamic host-feeding and oviposition behavior of an aphid parasitoid <Emphasis Type="Italic">Aphelinus asychis</Emphasis>

In order to maximize the lifetime reproductive success of parasitoids, they should be induced to dynamically accept individual hosts that have different suitability for oviposition. Parasitoids tend to exhibit higher host-selective behavior when their egg load is limited, and are less selective if they are facing time constraints. Here, we evaluated the effects of parasitoid age on egg load, fecundity and host instar preference of a honey-fed aphid parasitoid, Aphelinus asychis Walker (Hymenoptera: Aphelinidae). Host selective experiment was conducted to measure host-preference of honey-fed A. asychis females at different ages, using the second and fourth instars of the green peach aphid Myzus persicae as their hosts. The results showed that the choice of host-instar for oviposition was significantly influenced by the parasitoid age. Honey-fed parasitoids in the age groups of 1, 5, 10 and 20 days tended to parasitize predominantly second-instar aphids, whereas 15-days old parasitoids showed no significant preference of host instars. On the other hand, host-feeding preference was not affected by parasitoid age. Parasitoid females of all ages preferred younger aphids to older aphids. This result could help evaluate the effectiveness of A. asychis for biological control of M. persicae when they encountered mixed-instar aphids in the field. In addition, the results might be helpful in assessing the host killing effects of other host-feeding parasitoids.     

Within-patch foraging behaviour of the aphid parasitoidAphidius funebris: plant architecture, host behaviour, and individual variation

The influence of plant architecture, host colony size, and host colony structure on the foraging behaviour of the aphid parasitoidAphidius funebris Mackauer (Hymenoptera: Aphidiidae) was investigated using a factorial experimental design. The factorial design involved releasing individual parasitoid females in aphid colonies consisting of either 10 or 20 individuals ofUroleucon jaceae L. (Homoptera: Aphididae) of either only larval instar L3 or a mixture of host instars, both on unmanipulated plants and on plants that had all leaves adjacent to the colony removed. Interactions between the parasitoid and its host were recorded until the parasitoid had left the plant. The time females spent on the host plant and the number of eggs laid varied greatly among females. Host colony size significantly affected patch residence time and the number of contacts between parasitoids and aphids. Plant architecture influenced the time-budget of the parasitoids which used leaves adjacent to the aphid colony for attacking aphids. Female oviposition rate was higher on unmanipulated plants than on manipulated plants. No further significant treatment effects on patch residence time, the number of contacts, attacks or ovipositions were found. Oviposition success ofA. funebris was influenced by instar-specific host behaviour. Several rules-of-thumb proposed by foraging theory did not account for parasitoid patch-leaving behaviour.     

Functional responses of aphid parasitoids,Aphidius colemani (Hymenoptera: Braconidae) and Aphelinus asychis (Hymenoptera: Aphelinidae)

We evaluated the functional responses of two aphid parasitoids: Aphidius colemani on the green peach aphid Myzus persicae (Hemiptera: Aphididae), and Aphelinus asychis on M. persicae and the potato aphid, Macrosiphum euphorbiae (Hemiptera: Aphididae). Parasitoid oviposition occurred at host densities of 5, 10, 20, 30, 50, 80 or 100 aphids for A. colemani and 5, 10, 20, 30 or 50 aphids for A. asychis. More M. persicae were parasitized by A. colemani than by A. asychis at an aphid density of 50. Among the three types of functional response, type III best described the parasitoid response to the host densities both in A. colemani and A. asychis. The estimated handling time was shorter for A. colemani than for A. asychis (0.017 and 0.043 d, respectively). The proportion of aphids that were parasitized exhibited the same characteristic curve among the three host-parasitoid combinations: a wave form that appeared to be a composite of a decelerating (as in type II) response at low host density and an accelerating-and-decelerating (as in type III) response at medium to high host density. We hypothesize that the novel host species (and its host plant), density-dependent superparasitism, and/or density-dependent host-killing may have induced the modified type III response.     

Mating or ovipositing? A crucial decision in the life history of the cabbage aphid parasitoid Diaeretiella rapae (M’Intosh)

1. The reproductive fitness of a parasitoid depends on its mating and ovipositing success. Virgin haplodiploid females can reproduce, but produce only males, and may diminish fitness by producing more male offspring than required. Therefore, females must decide on whether to mate or oviposit first. 2. This study was conducted to assess the mating versus ovipositing decision and its impact on the reproductive fitness of Diaeretiella rapae (Hymenoptera: Aphididae), an endoparasitoid of the cabbage aphid Brevicoryne brassicae (Hemiptera: Aphididae). 3. When newly emerged females were given a choice between mating and ovipositing, about 62% of D. rapae females preferred to mate before ovipositing. Those females who oviposited before mating parasitised only 10% of the available aphids. After mating, females superparasitised their hosts with fertilised eggs, which resulted in a highly female‐biased sex ratio in the offspring. 4. Mating success was very high (91%) in the presence of hosts (cabbage aphid nymphs) compared with that in the absence of aphids. However, mating success was not influenced by the quality (size) of the hosts present in the mating arena, despite a parasitoid preference for larger hosts during oviposition. The time between pairing and mating was also shorter in the presence of host aphids. The mean number of aphids parasitised and the parasitism rate were significantly greater after mating.     

Larvicidal activity of lectins onLucilia cuprina: mechanism of action

Foraging behaviour and host-instar preference of young and old females of the solitary aphid parasitoid,Lysiphlebus cardui Marshall (Hymenoptera: Aphidiidae), were studied in the laboratory. The analysis of interactions between parasitoids and different stages ofAphis fabae cirsiiacanthoidis Scop. (Homoptera: Aphididae) revealed that encounter rates between aphids and parasitoid females and defence reactions of the aphids influenced the degree to which a particular aphid age class is parasitized. Encounter rates between hosts and parasitoid females depended on the foraging pattern of the parasitoid, which varied with age. In mixed aphid colonies patch residence time increased with parasitoid age. Furthermore, younger parasitoids (≦1 day old) laid more eggs into second and third instars, while older parasitoids (≧4 days old) did not show distinct host instar preferences. It is suggested that the oviposition behaviour ofL. cardui is influenced by the physiological state, i.e. the age of the wasp.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Cascading effects of N input on tritrophic (plant–aphid–parasitoid) interactions

Because N is frequently the most limiting mineral macronutrient for plants in terrestrial ecosystems, modulating N input may have ecological consequences through trophic levels. Thus, in agro‐ecosystems, the success of natural enemies may depend not only from their herbivorous hosts but also from the host plant whose qualities may be modulated by N input. We manipulated foliar N concentrations by providing to Camelina sativa plants three different nitrogen rates (control, optimal, and excessive). We examined how the altered host‐plant nutritional quality influenced the performances of two aphid species, the generalist green peach aphid, Myzus persicae, and the specialist cabbage aphid, Brevicoryne brassicae, and their common parasitoid Diaeretiella rapae. Both N inputs led to increased N concentrations in the plants but induced contrasted concentrations within aphid bodies depending on the species. Compared to the control, plant biomass increased when receiving the optimal N treatment but decreased under the excessive treatment. Performances of M. persicae improved under the optimal treatment compared to the control and excessive treatments whereas B. brassicae parameters declined following the excessive N treatment. In no‐choice trials, emergence rates of D. rapae developing in M. persicae were higher on both optimum and excessive N treatments, whereas they remained stable whatever the treatment when developing in B. brassicae. Size of emerging D. rapae females was positively affected by the treatment only when it developed in M. persicae on the excessive N treatment. This work showed that contrary to an optimal N treatment, when N was delivered in excess, plant suitability was reduced and consequently affected negatively aphid parameters. Surprisingly, these negative effects resulted in no or positive consequences on parasitoid parameters, suggesting a buffered effect at the third trophic level. Host N content, host suitability, and dietary specialization appear to be major factors explaining the functioning of our studied system.