Why is Coccidoxenoides perminutus, a mealybug parasitoid, ineffective as a biocontrol agent—Inaccurate measures of parasitism or low adult survival?

Coccidoxenoides perminutus achieves only low levels of parasitism of its host Planococcus citri in southeast Queensland citrus. Two possible causes were investigated. Adult survival under natural conditions was assessed to determine whether providing adult food sources could enhance survival. Behavioural changes of hosts, induced by C. perminutus parasitism, was also investigated to establish if parasitised P. citri move from their feeding site to seek protected shelters some distance away and are thus not accounted for in field assessments of parasitism rates. Unparasitised mealybugs placed in the field for two periods were retrieved before the effects of parasitism were manifested and parasitism rates were still low (0.3% at 5 days and 1.2% at 10 days). Levels of locomotion of P. citri exposed to C. perminutus were compared with those of “unexposed” ones. Parasitised mealybugs, regardless of instar, undergo behavioural changes. In comparison to unparasitised controls, the mealybugs become highly active 7–14 days after exposure to wasps. All parasitised mealybugs undergo physical changes, their body becomes cylindrical, their legs go so rigid that the mealybugs become immobile, and this signifies the typical mummy appearance. All mealybugs that became mummies eventually fell from the host lemon fruit because of impaired locomotion and were caught on sticky traps that had been placed beneath the lemons. Consequently, their final site of mummification was not established. C. perminutus adults provided with nectar or honey survived longer (about 5 days) in the field than those without food (about a day). Nectar from two plant species, Alpinia zerumbet and Datura candida, proved to be good sources of food for the adult wasps, and were comparable in quality to honey. The low level of parasitism achieved by C. perminutus in southeast Queensland citrus thus appears to be a consequence of the short adult life and the negative effects of a harsh environment. Provision of a suitable food source (e.g., nectar) may well enhance levels of parasitism in the field.     

Host location and discrimination mediated through olfactory stimuli in two species of Encyrtidae

The role of olfactory stimuli in host detection and evaluation was studied in two encyrtid Hymenoptera. The first, Epidinocarsis lopezi De Santis, is a monophagous parasitoid of the cassava mealybug Phenacoccus manihoti Matile-Ferrero, itself feeding exclusively on cassava, Manihot esculenta Crantz. The second, Leptomastix dactylopii Howard, is a monophagous parasitoid of the Citrus mealybug, Planococcus citri Risso, but this latter is highly polyphagous. The behaviour of females of both parasitoids (attaction and locomotion) was compared in a tubular olfactometer for the odours of their respective hosts on cassava and poinsettia. Tests were made using: 1) healthy host-plant alone; 2) host-plant infested with unparasitized mealybugs; 3) unparasitized mealybugs only; 4) host-plant infested with parasitized mealybugs and 5) parasitized mealybugs only. Only E. lopezi was attracted by the odour of the host-plant alone, but both species were attracted by the odour of an infested host-plant and that of unparasitized mealybugs. The odour of parasitized mealybugs, alone or on host-plant, induced an undirected activity. The attraction of E. lopezi to the odour of the host-plant alone could be linked to the monophagous diet of its host, whereas the attraction of the two species of parasitoids to the odours of infested host-plants and unparasitized mealybugs could be due to the fact that both parasitoids are specialists. The behavioural response of both species to the odour of parasitized mealybugs revealed a new aspect in host discrimination: the identification of parasitized hosts could be partly mediated through olfactory stimuli, and not only through gustatory stimuli.     

Larval parasitism of the autumnal moth reduces feeding intensity on the mountain birch

Plants respond to grazing by herbivorous insects by emitting a range of volatile organic compounds, which attract parasitoids to their insect hosts. However, a positive outcome for the host plant is a necessary precondition for making the attraction beneficial or even adaptive. Parasitoids benefit plants by killing herbivorous insects, thus reducing future herbivore pressure, but also by curtailing the feeding intensity of the still living, parasitised host. In this study, the effect of parasitism on food consumption of the 5th instar larvae of the autumnal moth (Epirrita autumnata) was examined under laboratory conditions. Daily food consumption, as well as the duration of the 5th instar, was measured for both parasitised and non-parasitised larvae. The results showed that parasitism by the solitary endoparasitoid Zele deceptor not only reduced leaf consumption significantly but also hastened the onset of pupation in autumnal moth larvae. On the basis of the results, an empirical model was derived to assess the affects on the scale of the whole tree. The model suggests that parasitoids might protect the tree from total defoliation at least at intermediate larval densities. Consequently, a potential for plant–parasitoid chemical signalling appears to exist, which seems to benefit the mountain birch (Betula pubescens ssp. czerepanovii) by reducing the overall intensity of herbivore defoliation due to parasitism by this hymenopteran parasitoid.     

Prey Habitat Location by the Cassava Mealybug Predator Exochomus flaviventris: Olfactory Responses to Odor of Plant, Mealybug, Plant–Mealybug Complex, and Plant–Mealybug–Natural Enemy Complex

Exochomus flaviventris Mader is considered to be the most active predator of the cassava mealybug Phenacoccus manihoti Matile–Ferrero in Central Africa. The response of experienced gravid female coccinellids to the odor of cassava plant (var. Zanaga), unparasitized mealybugs, plant–mealybug complex with or without feeding prey (parasitized or not), and plant–mealybug complex with or without conspecific coccinellids was investigated in a Y-tube olfactometer. The odor of uninfested cassava plants was not more attractive than clean air. Dual-choice tests revealed that mealybug-infested plants were preferred to mealybugs alone and mealybug-damaged plants and were the major sources of volatiles that attract females coccinellids to the microhabitat of its prey. The emission of volatile chemicals did not appear to be limited to the infested parts of the plant but did occur systemically throughout the plant. The presence of conspecific coccinellid larvae or adult males did not modify the attractiveness of the mealybug-infested plants. However, when an infested plant with conspecific predator females (alone or with conspecific males) was compared to an infested plant or infested plant with conspecific males, E. flaviventris females showed a preference for the last two sources of odor. The uninfested plant with conspecific males was also preferred to the uninfested plant with conspecific females. In addition, the odor of conspecific males was preferred over that of conspecific females. Female predators preferred the plant infested with unparasitized mealybugs over the plant infested with mealybugs previously parasitized. These results showed that E. flaviventris females use herbivore-induced plant volatiles during foraging and can detect via olfaction the presence of conspecific gravid females and parasitized prey, thus assessing patch suitability from a distance.     

The influence of two endoparasitic wasps, <Emphasis Type="Italic">Hyposoter didymator</Emphasis> and <Emphasis Type="Italic">Chelonus inanitus</Emphasis>, on the growth and food consumption of their host larva <Emphasis Type="Italic">Spodoptera littoralis</Emphasis>

The influence of parasitism by Hyposoter didymator (Thunberg; Hymenoptera: Ichneumonidae) and Chelonus inanitus (Linnaeus) (Hymenoptera: Braconidae) on the growth and food consumption of their host Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae) was studied in the laboratory. Parasitised larvae consumed significantly less artificial diet than unparasitised ones. Egg parasitisation by C. inanitus affected host larval consumption from the second day after emergence and it was significantly different from that of unparasitised ones. H. didymator, however, started to reduce larval consumption 4 days after parasitisation on the third instar host larvae. The overall reduction achieved by the larval endoparasitoid H. didymator is higher than that caused by the egg-larval endoparasitoid C. inanitus. The final body weight of a parasitised host larva by H. didymator and C. inanitus was only 6.7 and 13.0% of the maximum weight of an unparasitised sixth instar larva respectively. Moreover, parasitised larvae never reached the last instar. Results indicated that parasitised larvae might cause considerable less damage to the host plant than unparasitised ones.     

Electrically recorded feeding behaviour of cassava mealybug on host and non-host plants

The feeding behaviour of cassava mealybug (Phenacoccus manihoti) was analysed by the electrical penetration graph technique (EPG, DC-system) and by light microscopy. This study confirmed a typical phloem-feeding behaviour with an aphid-like predominance of extracellular pathways of stylets. Similarities of EPGs from mealybugs with those of aphids and whiteflies allowed adoption of standard pattern labelling. The main differences with aphid EPGs consisted of fewer but longer intracellular punctures (pd duration of 20 svs 5–7 s for aphids), longer times to the first phloem-ingestion period and a lower motility of stylets within the phloem searching process. Comparison of the feeding behaviour on 6 plant types, including two occasional hosts (Talinum and poinsettia), one cassava hybrid and three common cassava cultivars (Manihot esculenta), allowed differentiation of penetration profiles. Intracellular punctures were shorter on occasional hosts, on which phloem finding and subsequent ingestion were markedly delayed although a great variability within plants persisted. Chemical analysis of some secondary plant substances revealed that none of the plants tested contained detectable amounts of alkaloids, and that cyanides were restricted to true hosts (cassavas andManihot hybrid). Levels of total flavonoids did not differ between hosts and non-hosts, in contrast to those of phenolic acids. Preliminary correlations with EPG paterns are discussed.

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Résumé Le comportement alimentaire de la cochenille du manioc (Phenacoccus manihoti) a été étudié par électrographie de pénétration (EPG, analyse en courant continu) et par microscopie optique. Cette étude a permis de confirmer le caractère typiquement phloémophage de l'alimentation de cet Homoptère, présentant une large prédominance du trajet extracellulaire des stylets, comme cela est également observé chez les pucerons. Les similitudes des tracés EPG effectués sur cochenille et sur pucerons ou aleurodes ont permis d'adopter la même typologie du signal. Les principales différences avec les pucerons concernent le nombre moins important et la durée plus longue des ponctions intracellulaires (20 s en moyenne contre 5 à 7 s chez les pucerons), l'allongement du temps d'accès au phloème et une moins grande mobilité des stylets lors de la phase de recherche du phloème. La comparaison du comportement alimentaire sur 6 plantes, parmi lesquelles deux h?tes occasionnels (Talinum et poinsettia), un hybride et trois variétés courantes de manioc (Manihot esculenta), a permis de différencier plusieurs profils de pénétration. Les ponctions intracellulaires se sont révélées plus courtes sur h?tes occasionnels, qui induisent également, malgré la variabilité observée, un allongement net de la période de localisation du phloème et donc un retard à l'ingestion. Aucune des plantes testées ne contient d'alcalo?des, et les composés cyanés se restreignent au genreManihot, h?te naturel de la cochenille. Les flavono?des totaux ne permettent pas de distinguer plantes h?tes et non-h?tes, à la différence des acides phénoliques.

     

Sustained biological control of the cassava mealybugPhenacoccus manihoti [Hom.: Pseudococcidae] byEpidinocarsis lopezi [Hym.: Encyrtidae] in Nigeria

Following the successful introduction ofEpidinocarsis lopezi (De Santis) for biological control of the cassava mealybug (CM)Phenacoccus manihoti Mat.-Ferr. in southwestern Nigeria in 1981 and 1982, 11 groups of cassava fields were sampled every 2 weeks up to 1988 for impact assessment. After 1984, CM populations remained mostly below 10 per tip despite the presence of native hyperparasitoids, demonstrating the long-term success of biological control byE. lopezi in the region. Indigenous polyphagous coccinellids were found only during peak host densities, whereas the specificE. lopezi was common throughout the year. During some periods, percentage parasitism indicated delayed density dependence. Since 89% of all sampled cassava tips had no CM at all and the parasitisme is very mobile, parasitization rates were also calculated for individual infested tips (N=4,878). Parasitism increased slightly with host density on tips having between 1 and 10 CM of the 3^rd and 4^th instars, indicating positive density dependence. Such tips comprised 64% of all infested tips. At higher host densities, parasitism rates fell rapidly. The results are discussed in view of different theories on population regulation by biological control agents.      

Occurrence and activity of <Emphasis Type="Italic">Bemisia tabaci</Emphasis> parasitoids on cassava in different agro-ecologies in Uganda

Bemisia tabaci Gennadius (Homoptera: Aleyrodidae) is the vector of cassava mosaic geminiviruses that cause cassava mosaic disease (CMD), which in turn causes devastating yield losses. Surveys were conducted from October 2000 to November 2001 in four agro-ecologies in Uganda to enhance the understanding of parasitoid fauna and parasitism of B. tabaci in cassava fields. Such an understanding is an essential prerequisite for the development of biological control methods of B. tabaci to complement current CMD control practices. Parasitoid abundance and parasitism efficiency varied between locations and sampling dates within the locations; highest parasitoid densities were observed at Namulonge in the Lake Victoria crescent while the lowest was at Kalangala. In all locations, parasitism was mainly due to Encarsia sophia Dodd and Girault and Eretmocerus mundus Mercet (all Hymenoptera: Aphelinidae). Two occasionally observed species included Encarsia mineoi Viggiani (Hymenoptera: Aphelinidae), only observed at Namulonge, and blackhead Encarsia (Hymenoptera: Aphelinidae) observed at Bulisa, Namulonge and Lyantonde. Parasitism efficiency was highest at Bulisa (57.9%), but ranged from 40.2 to 46.9% at the other three sites. This paper discusses the possible causes of variations in parasitoid abundance and parasitism efficiency, and proposes further studies that might be carried out to assess the potential for augmentation of parasitoids to control B. tabacipopulations and CMD.     

Influence of water-stressed cassava on Phenacoccus herreni and three associated parasitoids

The influence of cassava Manihot esculenta Crantz grown under condition of water-stress on development and reproduction of the cassava mealybug, Phenacoccus herreni Cox & Williams, and levels of parasitism of three encyrtid parasitoids, Apoanagyrus diversicornis Howard, Aenasius vexans Kerrich, and Acerophagus coccois Smith, were studied in the laboratory. Two cassava cultivars were used: CM 507-37 (drought-tolerant) and CMC 40. A 30 day period of water stress, imposed by reducing the irrigation volume, led to a reduction in shoot development and stomatal conductance of leaves of both cassava genotypes. Phenacoccus herreni development and reproduction were favoured by cassava under water shortage. Parasitism decreased and water stress appeared to enhance the encapsulation of parasitoid eggs or larvae by the mealybug. In the case of the parasitoid A. diversicornis, there was a decrease in size of female progeny, suggesting a lower fitness in this species on cassava plants under water stress. All results indicated that cassava grown under low water availability favoured P. herreni development and reproduction, and affected the success of parasitism and, depending on the species, parasitoid development. The drought-tolerance characteristic of cassava genotypes and parasitoid species most suitable for controlling P. herreni in drought-stricken areas are discussed.     

Effect of Manipulations on the Host Selection Behavior of Sitobion avenae (Homoptera: Aphididae)

The study of aphid host selection and feeding behavior is difficult because aphids have to penetrate the plant to reach their feeding site, phloem tissue. The activity of the stylets, salivation or food intake, can not be observed externally and requires an indirect visualization technique such as the Electric Penetration Graph (EPG). The plant selection behavior of Sitobion avenae on potato varied depending on whether an ethological or EPG method was used to study it. A similar variation did not occur with Myzus persicae or Rhopalosiphum padi. The application of water-based silver conductive paint onto the thorax, as normally used for EPG, or onto the abdomen of Sitobion avenae alates resulted in increased duration and frequency of probing compared to results from ethological observations. Our results indicated that EPG manipulations might have different effects on different species of aphids and that a comparison of EPG and ethological data is required to confirm that the EPG method does not bias aphid feeding behavior.     

Interactions between the Parasitoid Rhopus nigroclavatus (Ashmead) (Hymenoptera: Encyrtidae) and its Mealybug Hosts Tridiscus sporoboli (Cockerell) and Trionymus sp. (Homoptera: Pseudococcidae)

This research investigated age-class-specific parasitism rates of the buffalograss mealybugs Tridiscus sporoboli (Cockerell) and Trionymus sp. by Rhopus nigroclavatus (Ashmead) (Hymenoptera; Encyrtidae), size class preference of this parasitoid, and mealybug–parasitoid interactions through choice and no-choice studies. In the no-choice studies, the mean rates of parasitism by R. nigroclavatus were 45, 20, 0, and 0%, respectively, for mealybugs adult female, third and fourth instars, first and second instars, and eggs. Choice studies indicated that rate of parasitism increased with host size. The mean rates of parasitism on mealybugs in the choice studies were 100% for adult females, 24% for third and fourth instars, 0% for first and second instars, and 0% for eggs. A second set of choice studies investigating mealybug/parasitoid behavior revealed that R. nigroclavatus oviposits in all post-egg mealybug age classes, but first and second instars were less often parasitized than older mealybugs.     

Vine mealybug sex pheromone increases citrus mealybug parasitism by Anagyrus sp. near pseudococci (Girault)

The present study was aimed at elucidating the role of lavandulyl senecioate (LS), the sex pheromone of Planococcus ficus, in host selection of the parasitoid Anagyrus sp. near pseudococci. Field trials were carried out in Portugal, Italy and Israel. The effect of LS on the parasitism rate of the wasp was determined by exposing sentinel mealybugs combined with pheromone dispensers impregnated with LS, in comparison with other baits: lavandulyl isovalerate (LI); planococcyl acetate (PAc); and unbaited control traps. In addition, in order to study the host location behavior of A. sp. near pseudococci, pheromone dispensers were placed at three different distances: inside the trap, 30 or 60 cm away from the trap. The number of parasitoid females inside the traps, the number of parasitized mealybugs, and the number of days required for the first parasitoid emergence were recorded. The response of A. sp. near pseudococci females to different doses of LS (25–1350 μg) was also evaluated using sticky plate traps. The rate of mealybug parasitism by A. sp. near pseudococci was significantly increased by LS in the three parasitoid populations. PAc and LI had no significant effect on the wasp parasitism rate in most of the trials. However, the Italian population of the parasitoid responded to PAc, showing apparently a different behavioral pattern. The number of parasitoid females trapped did not significantly differ between tested doses of LS. The use of LS as an arrestant in host location by the A. sp. near pseudococci female is discussed.     

Host specificity and comparative foraging behaviour of Aenasius vexans and Acerophagus coccois, two endo-parasitoids of the cassava mealybug

Two encyrtid parasitoids, Aenasius vexans Kerrich (Hymenoptera: Encyrtidae) and Acerophagus coccois Smith (Hymenoptera: Encyrtidae), were compared for their degree of dietary specialisation and the impact this has on their foraging strategies. Both parasitoid species are significant for biological control of the cassava mealybug, Phenacoccus herreni, Cox & Williams (Homoptera: Sternorrhyncha) a major Latin American pest of cassava, Manihot esculenta Crantz, an important root crop. Host acceptance and parasitism were analysed in seven mealybug species (with different levels of polyphagy) occurring in and around cassava fields. Results demonstrate that, in this ecosystem, An. vexans is a specialist for P. herreni while Ac. coccois is a generalist on the first and second trophic level. Of the seven mealybug species, P. herreni and P. madeirensis Green were the most acceptable hosts for Ac. coccois, followed by Ferrisia virgata Cockerell. Ac. coccois did not accept the other four mealybug species.The foraging and oviposition behaviour of individual parasitoids was observed in bioassays with cassava leaves infested by P. herreni. The two species used different strategies to locate their host. Aenasius vexans spent significantly more time walking and standing on an infested leaf and examined a host longer than did Ac. coccois. Acerophagus coccois, in contrast, spent more time for oviposition. As a consequence An. vexans parasitised more hosts in a given time than did Ac. coccois. Because the rate of offspring production of the two species did not differ, we conclude that the gregarious Ac. coccois's strategy to deposit several eggs at once might compensate for its relatively low number of ovipostitions, compared with the solitary An. vexans. These findings suggest that, given the advantages and limitations of each species, a multi-species approach to biological control of P. herreni may yield best results.     

Relative oviposition rates of the cassava hornworm,Erinnyis ello [lep.: Sphingidae], and accompanying parasitism byTelenomus sphingis [Hym.: Scelionidae], on upper and lower leaf surfaces of cassava

Oviposition by the cassava hornworm,Erinnyis ello L., was quantified for upper (abaxial) and lower (adaxial) leaf surfaces of cassava in the department of Tolima, Colombia. Relative rates of egg parasitism byTelenomus sphingis Ashmead on the different leaf surfaces were determined and compared. Approximately 6% of hornworm eggs had been placed on lower leaf surfaces. These eggs showed significantly lower rates of parasitism indicating reduced searching time or efficiency by the parasitoid on leaf undersides.      

Field placement of mealybug-infested potted cassava plants for the study of parasitism of Phenacoccus herreni

During 4-day field exposure intervals, Phenacoccus herreni Cox & Williams, third-instar mealybugs on artificially infested potted cassava plants were reduced 73% from initial densities of 235.7–1100.2/infested plant to 63.7–314.2/plant due to dispersal, rain, and predation. Frequency of parasitoid attack did not differ between high and low mealybug densities. The same species of parasitoids, in approximately the same proportions, were reared from mealybugs on artificially infested potted plants as from field mealybugs (Acerophagus coccois) Smith, 88.1% and Epidinocarsis diversicornis (Howard), 11.3%. Parasitoid attack rates on artificially exposed mealybugs, when combined with a simulation model of parasitism, accurately predicted observed levels of stage-specific parasitism in the field population.

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Résumé Pendant des expositions de 4 jours dans la nature de plants en pots de M. esculenta contaminés artificiellement par des P. herreni du troisième stade, les densités de cochenilles ont été réduites de 73%: de 235.7–1100.2/par plant contaminé à 63.7–314.2/par plant, par suite des évasions, de la pluie et de la prédation. Les attaques de parasitoïdes ne changent pas suivant les densités de cochenilles. Les mêmes espèces de parasitoïdes, ont été élevées dans à peu près les mêmes proportions (Acerophagus coccois, 88.1%, et Epidinocaris diversicornis, 11.3%) à partir de cochenilles sur pots artificiellement contaminés ou de manioc de la nature. Les taux d'attaques par les parasitoïdes sur les cochenilles artificiellement exposées, associés à un modèle de simulation du parasitisme, ont permis de prévoir avec précision les niveaux de parasitisme observés pour chaque stade dans la population de la nature.

     

The biology of two Eretmocerus spp. (Haldeman) and three Encarsia spp. Forster and their potential as biological control agents of Bemisia tabaci biotype B in Australia

The performance, as measured by daily rate of parasitism and total parasitism, of five aphelinid species found in Australia parasitising Bemisia tabaci were compared on cotton, hibiscus, rockmelon, soybean and tomato. Two Eretmocerusspp., both indigenous to Australia, gave the highest levels of parasitism on each of the plant host species tested. The tritrophic interactions between B. tabaci, host plant species and Eret. mundus(Australian parthenogenetic form) (APF) were also examined. In general, more whiteflies were parasitised when cotton was the source host or rockmelon the test host. Parasitism was always low when tomato was either the source or test host. When parasitoids were transferred from rockmelon to cotton, parasitism declined. In contrast, parasitism increased when parasitoids were transferred from cotton to rockmelon. Parasitism also increased when parasitoids were transferred from soybean to rockmelon, yet failed to do so when shifted from soybean to cotton despite cotton normally being a better host. However, when parasitoids were transferred from cotton to soybean there was a marked increase in parasitism. Possible causes are discussed. The field cage trial demonstrated that parasitism by both Eretmocerus spp. increased with increasing whitefly density. Further, the increase in parasitism was not due to the presence of more parasitoids as neither the parasitoid-whitefly ratio nor the total number of parasitoids present had a significant effect on parasitism. The combination of the two species, gave similar levels of parasitism to that achieved by Eret. mundus(APF) alone. Subsequent identification of the emerged individuals indicated that over 50% of the parasitism was due to this species suggesting that it out-competed Eret. queenslandensis. Despite this competition, there was no evidence that overall control was compromised.     

Assessing augmentative releases of parasitoids using the «Recruitment method», with reference toEdovum puttleri,a parasitoid of the colorado potato beetle [Coleoptera: chrysomelidae]

Release of approximately 17,700 experienced adult femaleEdovum puttleri Grissell against 1^st generation Colorado potato beetle eggs in 1987 in a 0.4 ha potato field in S. Deerfield, Massachusetts resulted in only 3.6% parasitism as assessed by direct measurement of host and parasitoid recruitment. Levels of non-viability indicated an additional 2.8% of hosts killed by parasitoid hostfeeding, for an overall impact of 6.4%. Release in 1988 of 126,300 parasitoid against 1^st generation hosts in a 0.4 ha potato (Solanum tuberosum L.) field at the same site produced only slightly higher levels of parasitism (10.6%) and host feeding (2.0%). Release in 1987 of 32,800 wasps against 2^nd generation eggs resulted in only 0.7% parasitism and 0.3% host feeding due to the toxicity of fenvalerate (Pydrin^R) residues from a single application applied for control of the potato leafhopper,Empoasca fabae (Harris). Release in 1988 of 47,400 wasps against the 2^nd host generation in the absence of any pesticide applications resulted in 34.4% parasitism and 16.1% host feeding, for a total impact of 50.5%. Difference in parasitization levels between host generations supports the idea thatE. puttleri adults require an in-field carbohydrate source such as aphid honeydew to reproduce. In Massachusetts, aphid populations in potato typically do not develop until the end of the 1^st larval generation. The recruitment method ofVan Driesche & Bellows (1988) proved to be a satisfactory approach for determining results of augmentative parasitoid releases.      

Intraguild predation of Planococcus ficus parasitoids Anagyrus pseudococci and Leptomastix dactylopii by Nephus kreissli

Intraguild predation of the mealybug parasitoids Anagyrus pseudococci (Girault), and Leptomastix dactylopii Howard (Hymenoptera: Encrytidae) by Nephus kreissli Fürsch & Uygun (Coleoptera: Coccinellidae) was studied. The latter is a native predator of the important pest Planococcus ficus (Signoret) (Hemiptera: Pseudococcidae) on grapevines in Turkey. For this purpose, P. ficus of different ages parasitised by A. pseudococci or L. dactylopii, or by both A. pseudococci and L. dactylopii, were served to fourth instars and adults of N. kreissli as food. Experiments were conducted using two different treatments: no-choice (served unparasitised or only one stage of parasitised mealybug) and choice (served unparasitised and only one age of parasitised mealybug together), under controlled environmental conditions. Both fourth instars and adult predators were fed on two- and four-day-old mealybugs parasitised by A. pseudococci or on two-, four- and six-day-old mealybugs parasitised by L. dactylopii or by either A. pseudococci or L. dactylopii. The predators could not consume six-day-old mealybugs parasitised by A. pseudococci, eight-day-old mealybugs parasitised by L. dactylopii, or those parasitised by either of these parasitoids which had become mummified. While it was found that the adult predators preferred parasitised mealybugs to unparasitised, the larvae did not show a pronounced preference between parasitised and unparasitised mealybugs (except for mealybugs parasitised by A. pseudococci).     

Altered dispersal behaviour in parasitised aphids: parasitoid-mediated or pathology?

1. Several hypotheses concerning modified dispersal behaviour in aphids parasitised by aphidiine wasps (Hymenoptera: Braconidae: Aphidiinae) were tested in the laboratory. Behavioural changes may be host-mediated, parasitoid-mediated, or a by-product of trauma and pathology. 2. Mummification site varied with parasitoid species. Pea aphids (Acyrthosiphon pisum) parasitised by Aphidius ervi, Aphidius pisivorus, Monoctonus paulensis, and Praon pequodorum mummified near the aphids’ preferred feeding sites on bean plants, but those parasitised by Ephedrus californicus often died and mummified outside the colony, away from the plants. 3. Parasitism by E. californicus had a progressive effect on the behaviour of pea aphids. Approaching death, aphids lost motor control and frequently dropped off the host plant when disturbed. Dropped aphids were unable to return to the feeding site and mummified elsewhere. The proportion of aphids mummifying outside the colony increased with mummy density. 4. Mummification site was not influenced by the presence within the same colony of aphids parasitised by different species of aphidiine wasps. 5. The evidence does not support the hypothesis that mummification site selection in E. californicus is determined by a host- or a parasitoid-mediated change in aphid dispersal behaviour. Association-specific differences in the dynamics of larval development and growth between aphidiine species provide an equally valid and possibly more general explanation of mummification behaviour.     

Floral diversity, parasitoids and hyperparasitoids – A laboratory approach

Adding floral resources to agro-ecosystems to improve biological control can enhance the survival, egg load, and parasitism rate of insect parasitoids. However, this may not always be the case because the herbivore may benefit from the added resource as much as, or more than the third-trophic level. In addition, the natural enemies of those in the third-trophic level may also derive improved fitness from the added resources. Both these processes will dampen trophic cascades, leading to less-effective biological control. In this study, the effect of adding different flowering plants on the longevity, egg load, aphid parasitism rates and hyperparasitism of Aphidius ervi Haliday (Hymenoptera: Braconidae) by its hyperparasitoid Dendrocerus aphidum Rondani (Hymenoptera: Megaspilidae) were investigated, using the pea aphid Acyrthosiphon pisum Harris (Homoptera: Aphididae) as the herbivore. Parasitoids exposed to buckwheat survived, on average, between four to five times as long as those in the control (water) and those in phacelia, alyssum and coriander treatments survived three to four times as long. Hyperparasitoids exposed to buckwheat survived five to six times as long as those in the control and three to five times longer with the other plants compared with the control. Almost all flower species significantly increased parasitoid and hyperparasitoid egg loads and the number of parasitised aphids and parasitised mummies compared with control. Understanding the factors influencing the dynamics of multitrophic interactions involving flowering plants, herbivores, parasitoids and hyperparasitoids is a fertile area for future research. One of the most challenging areas in contemporary ecology concerns the relative importance of different types of biodiversity mediating trophic interactions and thereby influencing the structure of communities and food webs. This paper begins to explore this using an experimental, laboratory-based approach.