Encapsulation and hemocyte numbers in three lepidopteran stemborers parasitized by Cotesia flavipes-complex endoparasitoids

Determination of the potential and actual host range of a natural enemy is crucial before its importation and release for biological control. We studied some of the factors that are important in determining the physiological host range of insect parasitoids attacking lepidopteran hosts. Our experimental system consisted of novel host-parasitoid associations, with two New World pyralid stalk borers, Diatraea saccharalis and D. grandiosella; one Old World crambid borer, Ostrinia nubilalis as hosts; and three Old World microgastrine braconids, Cotesia chilonis, C. sesamiae, and C. flavipes as parasitoids. Experiments on the chronology of encapsulation of the parasitoid progeny by host hemocytes indicated that lepidopteran stemborers that are taxonomically, behaviorally and ecologically very similar differ in their ability to encapsulate a parasitoid species. D. saccharalis encapsulated C. flavipes sometimes, whereas D. grandiosella consistently encapsulated C. sesamiae and C. flavipes. C. chilonis was not encapsulated by either Diatraea host. If encapsulation occurred it did not start until four days after parasitization and continued during the following days. O. nubilalis was an unsuitable host for all three parasitoid species; parasitoid eggs were killed within 24 hours of parasitization. O. nubilalis had nearly twice as many hemocytes present in the hemolymph compared to the Diatraea species. In many of the host-parasitoid combinations, there was an initial increase of hemocyte number soon after parasitization, which was not due to mechanical damage at oviposition. There was no correlation between total numbers of hemocytes present in the host hemolymph and the observed encapsulation levels. By understanding the encapsulation response we may be able to make better predictions about the host range of a parasitoid species before its release as a biological control agent.     

Host suitability of the New World stalkborer Diatraea considerata for three Old World Cotesia parasitoids

Biological control ofstalkboring Lepidoptera often has beensuccessful when the braconid parasitoids in thegenera Cotesia and Apanteles werethe natural enemies of choice. Constraints inusing these gregarious, koinobiont,endoparasitoids have included host suitability,especially as influenced by the host's immuneresponse. The suitability of a novel host, theNew World stalkborer Diatraeaconsiderata (Lepidoptera:Pyralidae), for parasitization by three OldWorld braconids, Cotesia chilonis, C. flavipes and C. sesamiae (Hymenoptera: Braconidae), wascompared to the suitability of another NewWorld novel host, Diatraea saccharalis. D. considerata was less suitable for allthree parasitoids than was D.saccharalis. The frequent occurrence ofparasitized D. considerata larvae thatdid not yield parasitoids, or pupate within anappropriate time interval, suggestedencapsulation of the parasitoid progeny, whichwas visible through the host cuticle. Giventhe suitability results, these threeparasitoids would not be appropriate candidatesfor use against D. considerata. Theresults also have important implications forthe narrow host range expressed by theseparasitoids.     

Development of the braconid wasp Cotesia flavipes in two Crambids, Diatraea saccharalis and Eoreuma loftini: Evidence of host developmental disruption

Cotesia flavipes is an important gregarious larval endoparasitoid of several crambid stem borers, including Diatraea saccharalis. The suitability of two crambid species, Eoreuma loftini and D. saccharalis, pests of sugarcane and rice in Texas, for C. flavipes development was tested. The effect of parasitization by C. flavipes on encapsulation response was assessed in vivo in both D. saccharalis and E. loftini. The results indicated that the parasitoid developed and emerged successfully in D. saccharalis larvae. Although E. loftini larvae were readily parasitized by C. flavipes parasitoids, no wasp larvae hatched from the eggs in this host because eggs were encapsulated by the host's hemocytes. The developmental fate of the E. loftini larvae with encapsulated parasitoids was variable. Most died as abnormal fifth instars or as post-wandering prepupae, while a few developed normally to the pupal stage. In vivo experiments, there was a significant reduction in the percent of beads encapsulated in parasitized larvae in both hosts. However, the percent of beads showing melanization decreased significantly in parasitized D. saccharalis larvae but did not differ significantly in parasitized or unparasitized E. loftini larvae. Our results showed that D. saccharalis is a suitable host for C. flavipes whereas E. loftini is an unsuitable host. This study indicated that lepidopteran stem borers that are taxonomically, behaviorally, and ecologically very similar can differ in their ability to encapsulate a parasitoid species.     

Evidences of non-additive effects of multiple parasitoids on Diatraea saccharalis Fabr. (Lep., Crambidae) populations in sugarcane fields in Brazil

Biological control is a relatively benign method of pest control. However, considerable debate exists over whether multiple natural enemies often interact to produce additive or non‐additive effects on their prey or host populations. Based on the large data set stored in the São João and Barra sugarcane mills (state of São Paulo, Brazil) regarding the programme of biological control of Diatraea saccharalis using the parasitoids Cotesia flavipes and tachinid flies, in the present study the author investigated whether the parasitoids released into sugarcane fields interfered significantly with the rate of parasitized D. saccharalis hosts. The author also observed whether there was an additive effect of releasing C. flavipes and tachinids on the rate of parasitized hosts, and looked for evidence of possible negative effects of the use of multiple parasitoid species in this biological control programme. Results showed that C. flavipes and the tachinids were concomitantly released in the Barra Mill, but not in the São Jão Mill. Furthermore, in the Barra Mill there was evidence that the parasitoids interacted because the percentage of parasitism did not increase after the release of either C. flavipes or tachinids. In the São João Mill, when both parasitoid species were released out of synchrony, both the percentage of parasitism by C. flavipes as well as that of the tachinids increased. When large numbers of tachinids were released in the Barra Mill, they caused a significant lower percentage of parasitism imposed by C. flavipes. The implications of the results as evidence of non‐additive effects of C. flavipes plus tachinids on D. saccharalis populations are discussed.     

Host Suitability of Four Cereal Stem Borers (Lepidoptera: Crambidae, Noctuidae) for Different Geographic Populations of Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) in Kenya

This study focused on the suitability of four species of cereal stem borers for the development of five geographic populations of Cotesia sesamiae (Cameron). C. sesamiae, an indigenous larval parasitoid of gramineous stem borers, is widespread in Africa. Four stem borers, Chilo partellus (Swinhoe), Chilo orichalcociliellus Strand (Lepidoptera: Crambidae), Busseola fusca Fuller, and Sesamia calamistis Hampson (Lepidoptera: Noctuidae), were offered to C. sesamiae for oviposition. Parasitoid individuals originated from five locations in Kenya. Biological parameters such as developmental time, percentage parasitism, progeny production, mortality of immature parasitoids, and proportion of female progeny were compared across host species. The two populations from western Kenya developed well on B. fusca. However, populations from the coast and the Eastern Province could not successfully parasitize B. fusca. With the exception of B. fusca, the percentage of hosts successfully parasitized by the different C. sesamiae populations was not different. The size of the host appeared to be an important factor influencing the development and reproductive potential of the parasitoid. We conclude that the different parasitoid populations were adapted to location-specific characteristics. Parasitoid–host compatibility must be evaluated before release for better establishment and colonization.     

Host–parasitoid community model and its potential application in biological control of cereal stemborers in Kenya

A two-host–two-parasitoid model was constructed to assess the effects of the introduced larval parasitoid, the braconid Cotesia flavipes, on its primary target host, the invasive crambid Chilo partellus, and on secondary host species, in inter-specific competition with Cotesia sesamiae, the main native parasitoid species of stemborers in Kenya. The model assumed that: (1) there was no host discrimination by either parasitoid species; (2) Cotesia flavipes was the superior competitor that out-competed Cotesia sesamiae when the host was suitable; and (3) Cotesia flavipes could only develop in an unsuitable host if it had been previously parasitized by Cotesia sesamiae. Model parameters were estimated from surveys conducted in Kenya and from laboratory experiments. Different scenarios of host and parasitoid species composition and host suitability occurring in the different ecological zones in Kenya were analyzed. Results indicated that: (1) the coexistence of stemborer host populations are determined by their population growth rates, the degree of aggregation of the parasitoids and their searching efficiency; (2) in the regions where both the invasive and the predominant native host species were suitable to either parasitoid species, stemborer densities would be reduced to and controlled at low densities, and Cotesia flavipes would become the dominant parasitoid species. However, the extinction or predominance of the native stemborer species depends on the ratio of the growth rates of exotic and native stemborers and their relative searching efficiencies; and (3) if the native host species was acceptable but unsuitable to Cotesia flavipes, the parasite would not become established.     

Immunosuppressive,antimicrobial and insecticidal activities of inhibitor cystine knot peptides produced by teratocytes of the endoparasitoid wasp Cotesia flavipes (Hymenoptera: Braconidae)

Teratocytes are specialized cells released by parasitoid wasps into their hosts. They are known for producing regulatory molecules that aid the development of immature parasitoids. We have recently reported the primary structures of cystine-rich peptides, including some containing inhibitor cystine knot (ICK) motifs, produced by teratocytes of the parasitoid Cotesia flavipes (Hymenoptera: Braconidae). ICKs are known for their stability and diverse biological functions. In this study, we produced four putative ICK peptides from the teratocytes of C. flavipes using solid-phase peptide synthesis or recombinant expression in E. coli, and investigated their functions on host immune modulation as well their potential to impair the development of two lepidopterans after ingestion of the peptides. In addition, the peptides were assayed against pathogens and human cells. The peptides did not influence total hemocyte count but suppressed cellular immunity, detectable as a reduction of hemocyte encapsulation (CftICK-I, CftICK-II, CftICK-III) and spread indexes (CftICK-IV) in the host. None of the peptides influenced the activities of prophenoloxidase and phenoloxidase in the hemolymph of larval Diatraea saccharalis (Lepidoptera: Crambidae). CftICK-I and CftICK-II with previously unknown function showed antifungal activity against Candida albicans but were non-toxic to human cells. CftICK-I, CftICK-II, and CftICK-III increased larval mortality and reduced leaf consumption of D. saccharalis, a permissive host for C. flavipes. The CftICK-III also increased larval mortality and reduced leaf consumption of Spodoptera frugiperda (Lepidoptera: Noctuidae), a non-permissive host for C. flavipes. This study highlights biological functions and biotechnological potential of ICK peptides from the teratocytes of C. flavipes.     

Sensilla on antennae,ovipositor and tarsi of the larval parasitoids,Cotesia sesamiae (Cameron 1906) and Cotesia flavipes Cameron 1891 (Hymenoptera: Braconidae): a comparative scanning electron microscopy study

Two braconid parasitoids of cereal stemborers in eastern Africa, Cotesia sesamiae and Cotesia flavipes, have been shown to display a similar hierarchy of behavioural events during host recognition and acceptance. In order to understand the mechanisms underlying host recognition and acceptance, the morphology of antennal sensilla on the last antennomeres, on the ovipositor, and on the fifth tarsomere and pretarsus of the prothoracic legs tarsi were studied using scanning electron microscopy followed by selective silver nitrate staining. It appeared that female C. sesamiae and C. flavipes shared the same types and distribution of sensory receptors, which enable them to detect volatiles and contact chemical stimuli from their hosts. In both parasitoids, four types of sensilla were identified on the three terminal antennomeres: (i) non-porous sensilla trichodea likely to be involved in mechanoreception, (ii) uniporous sensilla chaetica with porous tips that have gustatory functions, (iii) multiporous sensilla placodea, which are likely to have olfactory function, and (iv) sensilla coeloconica known to have thermo-hygroreceptive function. The tarsi of both parasitoids possessed a few uniporous sensilla chaetica with porous tips, which may have gustatory functions. The distal end of the ovipositor bore numerous dome-shaped sensilla. However, there were no sensilla coeloconica or styloconica, known to have gustatory function in other parasitoid species, on the ovipositors of the two braconid wasps.     

Foraging behavior and life history of the stemborer parasitoidCotesia flavipes (hymenoptera: Braconidae)

The gregarious parasitoidCotesia flavipes (Hymenoptera: Braconidae) attacks larvae of pyralid and noctuid stemborers by entering the stemborer tunnel. The short-range foraging behavior of femaleC. flavipes was studied on stemborerinfested plants, in patches with host-related products and in artificial transparent tunnels. In addition, the longevity under specific conditions and the potential and realized fecundity of femaleC. flavipes were determined. Larval frass, caterpillar regurgitate, and holes in the stem are used in host location byC. flavipes. The response to host products byC. flavipes seems not to be host species specific. FemaleC. flavipes respond to frass from four stemborer species and one leaf feeder. No differences are found in the behavior ofC. flavipes on maize plants infested with the suitable host,Chilo partellus (Lepidoptera: Pyralidae), or the unsuitable host,Busseola fusca (Lepidoptera: Noctuidae). Attacking a stemborer larva inside the stem is risky for the parasitoid. The mortality rate of the parasitoids inside the stem is high: 30–40% of the parasitoids are killed by the spitting and biting stemborer larva.C. flavipes is relatively shortlived: without food the parasitoids die within 2 days; with food and under high-humidity conditions they die within 5–6 days.C. flavipes is proovigenic and has about 150 eggs available for oviposition. A relatively large proportion of the available egg load (20–25%) is allocated to each host, so femaleC. flavipes are egg depleted after parasitizing only five or six hosts.     

Close-Range Host Searching Behavior of the Stemborer Parasitoids <Emphasis Type="Italic">Cotesia sesamiae</Emphasis> and <Emphasis Type="Italic">Dentichasmias busseolae</Emphasis>: Influence of a Non-Host Plant <Emphasis Type="Italic">Melinis minutiflora</Emphasis>

Studies were conducted on the host searching behavior of the larval parasitoid Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) and the pupal parasitoid Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae), both of which attack lepidopteran (Crambidae, Noctuidae) cereal stemborers. The behavior of D. busseolae was observed in a diversified habitat that consisted of stemborer host plants (maize, Zea mays L. and sorghum, Sorghum bicolor (L). Moench (Poaceae)) and a non-host plant (molasses grass, Melinis minutiflora Beauv. (Poaceae)), while C. sesamiae was observed separately on host plants and molasses grass. In previous olfactometer studies, C. sesamiae was attracted to molasses grass volatiles while hboxD. busseolae was repelled. The aim of the present study was to investigate the influence of molasses grass on close-range foraging behavior of the parasitoids in an arena that included infested and uninfested host plants. Dentichasmias busseolae strongly discriminated between host and non-host plants, with female wasps spending most of the time on infested host plants and least time on molasses grass. Likewise, C. sesamiae spent more time on uninfested and infested host plants than it did on molasses grass in single choice bioassays. While on infested plants, the wasps spent more time foraging on the stem, the site of damage, than on other areas of the plant. Overall, the results indicate that presence of the non-host plant does not hinder close range foraging activities of either parasitoid.     

Wolbachia infection dynamics in experimental laboratory populations of Metaseiulus occidentalis

The present study examined the acceptability and suitability of Old World stemborers (Chilo partellus and C. orichalcociliellus) for the development of New World parasitoids (Apanteles deplanatus and A. minator) and New World stemborers (Diatraea saccharalis and D. grandiosella) for the development of Old World parasitoids (Cotesia sesamiae, C. flavipes and C. chilonis). Results revealed that acceptance and suitability were high in old associations. In new associations, parasitoids accepted about 60% of the new association hosts. In addition, 10 out of 17 new associations were successful. Apanteles species appeared to be more physiologically host specific than Cotesia species. For example, two of four new association hosts were accepted by A. deplanatus and only one (D. saccharalis) was partially suitable for progeny development. Among the Cotesia species, Cotesia flavipes appeared to have a wider host range than the two other species. It attacked all hosts offered and successfully parasitized all but one (D. grandiosella). Diatraea saccharalis was accepted and was a suitable host for the development of all parasitoid species tested, whereas D. grandiosella was unsuitable for the development of four out of five parasitoid species tested. No clear pattern was observed as behavioral acceptance did not always agree with the pattern of physiological suitability. Implications of these findings for importation biological control of stemborers are discussed.     

Effects of single and multiple herbivory by host and non‐host caterpillars on the attractiveness of herbivore‐induced volatiles of sugarcane to the generalist parasitoid Cotesia flavipes

It is well known that parasitoids are attracted to volatiles emitted by host‐damaged plants; however, this tritrophic interaction may change if plants are attacked by more than one herbivore species. The larval parasitoid Cotesia flavipesCameron (Hymenoptera: Braconidae) has been used intensively in Brazil to control the sugarcane borer, Diatraea saccharalisFabricius (Lepidoptera: Pyralidae) in sugarcane crops, where Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), a non‐stemborer lepidopteran, is also a pest. Here, we investigated the ability of C. flavipes to discriminate between an unsuitable host (S. frugiperda) and a suitable host (D. saccharalis) based on herbivore‐induced plant volatiles (HIPVs) emitted by sugarcane, and whether multiple herbivory (D. saccharalis feeding on stalk + S. frugiperda feeding on leaves) in sugarcane affected the attractiveness of HIPVs to C. flavipes. Olfactometer assays indicated that volatiles of host and non‐host‐damaged plants were attractive to C. flavipes. Even though host‐ and non‐host‐damaged plants emitted considerably different volatile blends, neither naïve nor experienced wasps discriminated suitable and unsuitable hosts by means of HIPVs emitted by sugarcane. With regard to multiple herbivory, wasps innately preferred the odor blend emitted by sugarcane upon non‐host + host herbivory over host‐only damaged plants. Multiple herbivory caused a suppression of some volatiles relative to non‐host‐damaged sugarcane that may have resulted from the unaltered levels of jasmonic acid in host‐damaged plants, or from reduced palatability of host‐damaged plants to S. frugiperda. In conclusion, our study showed that C. flavipes responds to a wide range of plant volatile blends, and does not discriminate host from non‐host and non‐stemborer caterpillars based on HIPVs emitted from sugarcane. Moreover, we showed that multiple herbivory by the sugarcane borer and fall armyworm increases the attractiveness of sugarcane plants to the parasitoids.     

Importance of contact chemical cues in host recognition and acceptance by the braconid larval endoparasitoids Cotesia sesamiae and Cotesia flavipes

The ability of the congeneric braconid parasitoids Cotesia sesamiae (Cameron) and Cotesia flavipes Cameron to discriminate between stemborer larval cues upon contact was studied using their natural hosts, namely the noctuid Busseola fusca (Fuller) and the crambid Chilo partellus (Swinhoe), respectively, and the pyralid non-host Eldana saccharina (Walker). When the natural host larvae were washed in distilled water, parasitoid behavior was similar to that displayed when in contact with E. saccharina, characterized by the absence of ovipositor insertion. When washed host or non-host larvae were bathed with water extracts of their natural host, the parasitoids showed a significant increase in ovipositor insertions. However, the water extracts of host-larvae deposited on cotton wool balls did not induce ovipositor insertion in either C. sesamiae or C. flavipes. Nevertheless, the extracts enabled the parasitoids to discriminate between natural and non-hosts as indicated by the intensive antennating of the former. For both parasitoids, frass was found to be important in short-range host recognition as indicated by differences in the time spent on antennating between frass sources. In addition, the regurgitants of B. fusca and C. partellus induced ovipositor insertion in C. flavipes only. These results indicated that C. sesamiae and C. flavipes used different chemical cues for acceptation and oviposition in a stemborer larva, and that B. fusca and C. partellus shared the same chemical cues to induce oviposition in C. flavipes.     

Longevity of multiple species of tephritid (Diptera) fruit fly parasitoids (Hymenoptera: Braconidae: Opiinae) provided exotic and sympatric-fruit based diets

While adult parasitic Hymenoptera in general feed on floral and extrafloral nectars, hemipteran-honeydews and fluids from punctured hosts, Diachasmimorpha longicaudata (Ashmead), an Old World opiine braconid introduced to tropical/subtropical America for the biological control of Anastrepha spp. (Tephritidae), can survive on fruit juices as they seep from injured fruit. An ability to exploit fruit juice would allow such a parasitoid to efficiently forage for hosts and food sources simultaneously. Two New World opiines, Doryctobracon areolatus (Szepligeti) and Utetes anastrephae (Viereck), are also prominent Anastrepha parasitoids and are roughly sympatric. All three species were provided with: (1) pulp and juice diets derived from a highly domesticated Old World fruit (orange, Citrus sinensis L.) that is only recently sympatric with the Mexican flies and parasitoids and so offered little opportunity for the evolution of feeding-adaptations and (2) a less-domesticated New World fruit (guava, Psidium guajava L.), sympatric over evolutionary time with D. areolatus and U. anastrephae. Both sexes of D. longicaudata died when provided guava pulp or juice at a rate similar to a water-only control. D. areolatus and U. anastrephae, presumably adapted to the nutrient/chemical constituents of guava, also died at a similar rate. Survival of all three species on orange pulp and juice was greater than on water, and often equaled that obtained on a honey and water solution. In confirmatory experiments in Mexico, D. areolatus and U. anastrephae, as well as other tephritid parasitoids Doryctobracon crawfordi (Viereck) and Opius hirtus (Fisher), all died at a significantly higher rates when provided guava in comparison to a honey and water diet. Such a result is likely due to guavas being repellent, innutritious or toxic. D. longicaudata clearly consumed guava juice tagged with a colored dye. Dilutions of orange and guava juice resulted in shorter lifespans than dilutions of orange juice and water demonstrating that there while diluted orange juice provided nutrition the addition of guava created toxicity. Given the differences in fruit-food quality, adult opiine food sources would not be obtainable at all oviposition sites and in the case of guava, more additional sites and foraging for food than previously postulated may be required.     

Host-plant mediated effects of transgenic maize on the insect parasitoid Campoletis sonorensis (Hymenoptera: Ichneumonidae)

Determining the impact of genetically modified (GM) crops on beneficial organisms is an important aspect of the environmental risk assessment of GM crops. In the present study, the impact of Bt maize expressing Cry1Ab on the development and behaviour of the parasitoid Campoletis sonorensis was compared to individuals reared on hosts fed conventionally bred plants partially resistant to the European corn borer (Ostrinia nubilalis Hübner) and on susceptible maize hybrids. Adult parasitoids reared on Bt maize-fed Spodoptera frugiperda larvae were significantly smaller (15–30%) than those reared in hosts fed either of the conventional maize hybrids. The magnitude of this effect was dependent on the size of the host at oviposition and its subsequent growth rate. The development time of C. sonorensis was not affected by the maize treatment. In choice tests, female parasitoids displayed no preference for hosts fed a specific maize hybrid. No Cry1Ab was detected within adult parasitoids.     

Benefits of self-superparasitism in a polyembryonic parasitoid

Macrocentrus grandii is a polyembryonic parasitoid, with embryos that divide asexually within the host (European corn borer, Ostrinia nubilalis) to produce broods of clonal offspring. From a biological control standpoint, polyembryony seems advantageous because each parasitized host yields multiple parasitoids with minimal time and egg investment. When we observed M. grandii in the field, however, we found that the parasitoid virtually always invested additional time and, if possible, stings into hosts that it had already stung, apparently reducing some of the advantages of polyembryony. We therefore investigated and found support for two potential benefits that can be gained by self-superparasitism in this system. First, multiple stings allowed production of mixed-sex broods: 27% of multiply-stung versus 0% of singly-stung hosts produced mixed-sex broods. Second, multiple stings increased mean parasitoid progeny produced per host, primarily by reducing the chance of complete brood failure. Our results indicate substantial benefit for a second sting, but little benefit for three or more stings, even though M. grandii was sometimes observed to invest more than two stings within a single host. However, we also found that within-host larval competition is prevalent, suggesting that supernumerary stings may pay off in competition against conspecific larvae. Such additional investment within a single host would be particularly beneficial when hosts, rather than eggs, are limiting, but would decrease the overall efficacy of M. grandii as a biological control agent.     

Consequences of mixed species infestation on the searching behavior and parasitism success of a larval parasitoid

When two herbivore pest species are potential hosts of a single parasitoid species, two questions arise. Firstly, which host is preferable for mass rearing in terms of later parasitoid performance, and secondly, how do parasitoids perform in mixed herbivore situations after colony establishment? We tested Hyssopus pallidus, a gregarious parasitoid of two major pests of apple, Cydia (Grapholita) molesta and Cydia pomonella, before and after landing on apples infested by one of the two Cydia species. Pre-alighting host preference was tested in a Y-tube olfactometer setup, and parasitism success in a contact bioassay. To gain information on parasitoid performance throughout the growing season, different fruit growth stages were used. Irrespective of the host they had developed on, the parasitoids showed similar olfactory preferences when given a dual choice between infested and healthy fruits, and they did not discriminate between fruits infested by C. molesta and C. pomonella. Responsiveness was generally high, especially late in the season close to harvest. Both hosts are parasitized regardless of the host the parasitoid female had developed on, and no differences in parasitism rates or number of offspring were noted for the two hosts offered. Results were consistent for all apple growth stages tested. In conclusion, mass rearing of this parasitoid can be carried out on either host, without limiting the future efficacy of the bio-control agent. Similarly, established colonies are expected to develop further on both hosts without any bias in host preference.     

Assessment ofTrichogramma species for biological control of forest lepidopteran defoliators

In a laboratory study, we determined the potential of threeTrichogramma (Hymenoptera: Trichogrammatidae) species,T. brassicae Bezdenko,T. minutum Riley andT. nr.sibiricum Sorokina, for biological control against six species of forest lepidopteran pests, black army cutworm, hemlock looper, eastern spruce budworm, western spruce budworm, white-marked tussock moth, and gypsy moth. Females of each parasitoid species were offered eggs from each of the six host species. Parasitization and the effect of the host species on the emerging progeny were examined and recorded.Trichogramma minutum had the broadest host range and successfully parasitized four host species out of the six offered.Trichogramma nr.sibiricum had the narrowest host range and parasitized only two species of hosts. Of the six host species, black army cutworm was the most preferred by all threeTrichogramma species; white-marked tussock moth and gypsy moth were not parasitized by any parasitoids. There was a positive correlation between the size of female offspring and their corresponding egg complement in all three parasitoid species. The developmental time of parasitoids from egg to adult was influenced by both the parasitoid and host species. Our results suggest thatT. minutum has the greatest potential for biological control against various forest lepidopteran pests and that the black army cutworm may be the best target candidate for further study.     

Dispersal of the exotic parasitoid Cotesia flavipes in a new ecosystem

A study on the dispersal of the exotic larval endoparasitoid, Cotesia flavipes Cameron (Hymenoptera: Braconidae), was conducted in a maize field in the northern Kilifi District in the coastal area of Kenya. Because C. flavipes did not previously occur in the release area, it was possible to use a unique indirect method to estimate dispersal by examining the distribution of parasitised hosts. Parasitoids released in the centre of the field moved as far as 64 meters during their life span, and dispersal was dependent on wind direction. The level of parasitism was influenced by the location of hosts in plants. The majority of parasitised stemborers (88.4%) were found inside the plant (stems and tassel stems), where 74.3% of the suitable hosts were found, which indicates that female parasitoids were not searching randomly for hosts. Aggregation of parasitoids in response to plants with different host densities was not detected. Implications of the release of C. flavipes on stemborers population in the agroecosystem of East Africa are discussed.     

The Bionomics ofCoptera Haywardi(Ogloblin) (Hymenoptera: Diapriidae) and Other Pupal Parasitoids of Tephritid Fruit Flies (Diptera)

The endoparasitoidCoptera haywardi(Ogloblin) (Diapriidae) was discovered in Mexico attacking the pupae of the Mexican fruit fly,Anastrepha ludens(Loew). Typically, parasitoids of Diptera Cychlorrhapha pupae develop as ectoparasitoids and are generalists that attack hosts in a number of families. Aspects of the bionomics ofC. haywardiwere compared to those of two chalcidoid ectoparasitoids,Dirhinus himalayanusWestwood andSpalangia geminaBoucek.C. haywardideveloped in three genera of Tephritidae, but not in species of other families. The two species of chalcidoids developed in all the calypterate and acalypterate hosts to which they were exposed. In an olfactometerC. haywardipreferredAnastrepha suspensa(Loew) pupae, while the chalcidoids preferred the pupae ofMusca domestica L.This preference inS. geminawas diminished in insects that had been reared onA. suspensa. C. haywardioviposited in theA. suspensapupae that had been previously parasitized by the braconidDiachasmimorpha longicaudata(Ashmead). However, it completed development only in unparasitized pupae. Mortality of the primary parasitoid due toD. himalayanuswas approximately two-thirds the mortality inflicted on the host fly.S. geminadid not discriminate between parasitized and unparasitized pupae ofA. suspensaand developed in both.C. haywardiappears to have a more restricted host range relative to chalcidoid pupal parasitoids and this may be due to its endoparasitic development.