Behavior and Activity Pattern of Cephalonomia stephanoderis (Hymenoptera: Bethylidae) Attacking the Coffee Berry Borer,Hypothenemus hampei (Coleoptera: Scolytidae)

We describe behavioral sequences and daily activities of pre-ovipositing and ovipositing females of Cephalonomia stephanoderis (Hymenoptera: Bethylidae), an ectoparasitoid of the coffee berry borer, Hypothenemus hampei (Coleoptera: Scolytidae). Noticeable behavioral differences among preovipositing and ovipositing females include host examination, host stinging—probing, host feeding, and the oviposition per se. The female of C. stephanoderis feeds primarily on host eggs, but pupae are also exploited, mainly by pre-ovipositing females. After the onset of the oviposition period, C. stephanoderis examines the pupae repeatedly, stings them at frequent intervals, and spends more time feeding than during the pre-oviposition period. Host paralysis is linked both to host feeding and oviposition. It induces irreversible developmental arrest, which presumably allows preservation of the host until subsequent utilization, and contributes to successful offspring development, particularly by reducing host motility. Oviposition consists in a host selection process, a prolonged period of preparation of the potential host, and the egg-laying phase itself. Under our experimental conditions, pre-ovipositing and ovipositing females are active 17% and 36% of the day, respectively. Host handling time averages 6% and 23% in pre-ovipositing and ovipositing females, respectively. All coffee berry borer developmental stages are exploited by C. stephanoderis females, either for host feeding and/or oviposition activities. Such flexible behavior is advantageous given that host availability is limited inside the coffee berries.     

Alternative Hosts for Bethylid Parasitoids of the Coffee Berry Borer, Hypothenemus hampei (Coleoptera: Scolytidae)

Three species of bethylid wasps, Prorops nasuta Waterston, Cephalonomia stephanoderis Betrem, and C. hyalinipennis Ashmead, attack the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae), by both predation and parasitism. C. hyalinipennis has only recently been reported to attack H. hampei. Its previously recorded hosts belong to the coleopteran families Curculionidae, Anobiidae, and Scolytidae. We evaluated five further coleopteran species, in the families Curculionidae, Bostrichidae, and Bruchidae, as alternative hosts. High proportions of all five species were both fed upon and oviposited on (normally by concurrent host feeding). Offspring developmental mortality varied greatly (12–95%) on different host species. Offspring production per host was about three times greater than on H. hampei when females were presented with the curculionids Caulophilus oryzae (Gyllenhal) and two species of Sitophilus. C. stephanoderis and P. nasuta are currently the main biological control agents of H. hampei: they have long been thought to be monophagous but C. stephanoderis has recently been reported to attack H. obscurus. We evaluated C. oryzae, as a potential host or prey for P. nasuta, and both C. oryzae and Sitophilus sp. for C. stephanoderis. Host feeding was very common in both parasitoid species. The majority of females also oviposited on C. oryzae (71% P. nasuta, 60% C. stephanoderis) and 20% of C. stephanoderis parasitized Sitophilus sp. Fifty-two percent of C. stephanoderis and 7% of P. nasuta offspring survived to adulthood when developing on C. oryzae. The lifetime fecundities of these parasitoids presented with these host species were, however, low (maximum 17 eggs, mean ≤ 8.25) and preoviposition periods were generally long (ca. 19 days). Adult female C. stephanoderis lived longer when presented C. oryzae rather than Sitophilus sp. Host feeding also increased longevity. We discuss these results in the context of mass rearing these bethylids for release as biological control agents.     

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.     

Survival of the parasitoidEncarsia formosa after treatment of parasitized greenhouse whitefly larvae with fungal spores ofAschersonia aleyrodis

The interaction between the entomopathogenic fungusAschersonia aleyrodis and the parasitoidEncarsia formosa on greenhouse whitefly as a host organism was studied, in particular, the survival of the parasitoid after treatment of parasitized hosts with fungal spores. The mean number of parasitized black pupae per parasitoid produced at 25°C was significantly reduced after spore treatment in the first three days following parasitization. Spore treatment four, seven or ten days after parasitization resulted in a mean number of parasitized pupae not significantly different from the number of black pupae in the control. The rather sudden change from low to high survival of parasitized hosts when treated with spores four days after parasitization in spite of high numbers of infected unparasitized larvae, coincided with the hatching of the parasitoid larva from the egg inside the host. Possible reasons for this decrease in susceptibility to infection after parasitoid egg hatch, such as induced changes in host cuticle or haemolymph, are discussed. Parasitoids emerged from treated hosts did not show differences in reproduction compared with parasitoids emerging from untreated hosts. Both natural enemeies of whitefly are compatible to a great extent.     

Intra- and interspecific host discrimination in two closely related egg parasitoids

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.     

Host-finding behavior, host acceptance, and host suitability of the parasiteXanthopimapla stemmator

Xanthopimpla stemmator (Thunberg)(Hymenoptera: Ichneumonidae), a solitary endoparasite of pupae of Old World lepidopteran stalkborers, was recently imported into Texas as a candidate for biological control of New World stalkborers. Information on host acceptability, host suitability and cues responsible for host finding were necessary to gain an insight into parasite/host interactions, because of the absence of a coevolutionary history.Xanthopimpla stemmator females were exposed to laboratory-reared one-to six-day-oldDiatraea saccharalis (F.) pupae. An average of 62% of host pupae were accepted and all ages of pupae were equally acceptable. Host suitability decreased with host age. One- to five-day-old host pupae averaged 31–37% suitability, whereas only 19% of 6-day-old pupae were suitable. Successful parasitization, defined as the product of the proportion accepted and the proportion suitable, decreased from 22–23% for 1-, 2- and 3-day-old pupae to 13% for 6-day-old pupae. Sex ratio (female:male) of the parasite progeny increased with host age. Females comprised 47% of total parasite progeny of 1-day-old and 84% of 6-day-old pupae. The increase in percent females was a result of a similar number of females in all age classes, coupled with a decrease in the number of males from older hosts.Xanthopimpla stemmator superparasitized 61% of acceptedD. saccharalis pupae in the laboratory. On dissection, 73% of host pupae with multiple probe wounds were found to contain parasite eggs or larvae; these hosts contained up to 10 eggs or 7 first-instar larvae. Increased numbers of probes by the parasites were associated with an increase in successful parasitization. Host seeking activity inX. stemmator was stimulated by the presence of larval frass, host odor and movement of host pupae. Results suggest thatX. stemmator is a good candidate for biological control ofD. saccharalis and possibly other factitious stalkborer hosts.     

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.     

Rearing phonotactic parasitoid flies [Diptera: Tachinidae,ormiini, ormia spp.]

Females of ormiine tachinids fly to their hosts' calling songs and deposit larvae on the host or nearby. Two species,Ormia ochracea (Bigot) andO. depleta (Wiedemann), were reared for at least 8 generations, making them the first ormiines to be laboratory-propagated. Both were reared on natural hosts:Gryllus spp. field crickets (principallyG. rubens) forO. ochracea, andScapteriscus spp. mole crickets forO. depleta. Commercially rearedAcheta domesticus tested as hosts were less satisfactory. Hosts were parasitized manually or by confinement with flies or planidia (infective larvae). Transparent, cylindrical, sleeved cages were designed to accommodate parasitized hosts and pupae and adults ofO. ochracea. Cages were joined to allowO. ochracea to cycle through its stages with minimum handling and care. Parasitized hosts and pupae ofO. depleta were held in containers of damp sand; adults were held in cages developed forO. ochracea. Adults of both species were maintained on applesauce, sugar cubes, powdered milk, and water. The life cycle ofO. ochracea was about 31 days and ofO. depleta about 36 days, with the principal difference being the time required for planidia to complete development. InO. ochracea the adults emerged synchronously but inO. depleta males preceded females. In both species sex ratio was generally I: 1 and females lived slightly longer than males.O. depleta from our laboratory colony have been released for biological control of mole crickets.      

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4-day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae-larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

Parasitization of the cottony-cushion scale in relation to host size

Measurements of body length of cottony-cushion scales,Icerya purchasi Maskell, are presented. Although length increased markedly with developmental stage, the length distributions of successive stages were found to overlap, making length an imperfect indicator of stage andvice versa. The likelihood of parasitism by the fly,Cryptochaetum iceryae (Williston), was found to increase with increasing scale size when scales of different sizes were concurrently made available to the parasites under field conditions. Also, parasite loads (no. of parasites per parasitized host) were found to increase with host size. The size ofC. iceryae pupae was found to depend on the developmental stage of the scale host in which pupation took place — the more developed (larger) the host, the larger the pupa. This result suggests that parasite growth is food limited in the smaller hosts, and that therefore its apparent preference for larger hosts is to the parasite's advantage.      

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

Egg allocation of the autoparasitoid encarsia tricolor at different relative densities of the primary host (Trialeurodes vaporariorum) and two secondary hosts (Encarsia formosa and E. tricolor)

Parasitism, offspring sex ratio and superparasitism of the facultative autoparasitoid Encarsia tricolor Foërster (Hymenoptera: Aphelinidae) when given access to arenae with different proportions of the primary host (Trialeurodes vaporariorum (Westwood)) and two species of secondary hosts (E. tricolor and Encarsia formosa Gahan) were studied.Parasitism and offspring sex ratio were not affected by female age in the range 3–10 days old. When the secondary hosts were young E. tricolor pupae, eggs were mostly laid on primary hosts, so the offspring sex ratio was more female-biased than expected, and secondary hosts were not superparasitized at all. When the secondary hosts were fully grown E. formosa larvae, superparasitism was small and offspring sex ratio was more male-biased than expected. E. tricolor females were able to discriminate between hosts previously parasitized by themselves and non-parasitized hosts.     

The effect of host size on quality attributes of the egg parasitoid,Trichogramma pretiosum

In a study of the quality ofTrichogramma pretiosum Riley (Hymenoptera: Trichogrammidae), we compared female wasps emerging from natural hosts, parasitized in the laboratory or the field with those emerging from factitious hosts used for commercial mass production. Females from the natural hosts were larger, more fecund, and longer lived than those from the factitious hosts. Compared to small females, large female wasps are substantially more fecund when honey (carbohydrate) is available but marginally more fecund when honey is unavailable. The size of a femaleT. pretiosum depends on two factors: the size of the host egg from which it emerges even when the wasp was gregarious, and the number of conspecifics that emerge with it. The similarities in the size distribution of female wasps emerging from natural hosts, in conjunction with the mechanism by whichTrichogramma measure host size and allocate eggs accordingly, suggests the hypothesis that size related components of fitness in femaleT. pretiosum are under strong selection in the field.     

Host-discrimination by <Emphasis Type="Italic">Phymastichus coffea</Emphasis>, a parasitoid of the coffee berry borer

Phymastichus coffea (LaSalle) (Hymenoptera: Eulophidae) is an African endoparasitoid of the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae) that has been introduced to several countries to control this important pest. In the present study we performed a series of laboratory experiments in order to determine if there was evidence of host discrimination and superparasitism in P. coffea. Our choice experiments demonstrate that P. coffea females showed significant preference to attack unparasitized hosts, rather than those parasitized conspecifically. No significant preferences were detected in self-specific attacks between parasitized hosts and the healthy ones. A further dissection of hosts sequentially attacked either self or conspecifically, revealed that there were no more than two eggs per host. As P. coffea is a species that normally allocates two eggs per host in a single attack, we assumed that females were able to attack already parasitized hosts, but they did not lay eggs in them. Based on this fact, we conclude that there is a host discrimination ability in P. coffea females. With respect to the superparasitism by P. coffea using non-choice experiments, there was no significant difference between self-specific or conspecific attacks with respect to the control after one or two successive attacks. Conspecific attacks yielded the largest numbers of eggs after 3rd, 4th and 5th attacks and significant differences were found between this treatment and the control. The maximum number of eggs found in a single host was six individuals (conspecific treatment). These results confirmed that P. coffea usually laid two eggs per host; however, when there are no hosts available, conspecific attacks can result in the superparasitism in this species.     

The parasitoidOoencyrtus nezarae (Hymenoptera: Encyrtidae) prefers hosts parasitized by conspecifics over unparasitized hosts

Summary Two laboratory experiments were conducted to examine the ovipositional preferences of the egg parasitoidOoencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) for parasitized and unparasitizedMegacopta punctatissimum Montandon (Hemiptera: Plataspidae). Females that had never oviposited or that had not oviposite for 3 days preferred recently parasitized hosts more than unparasitized hosts. The preference for recently parasitized hosts appeared to be mediated by the punctures in already parasitized hosts made by the ovipositor of the first female. Survival of the parasitoid progeny was lower in recently parasitized hosts than in unparasitized hosts. However, handling time of parasitized hosts was extremely short relative to that of unparasitized hosts, because the superparasitizing female could use the punctures made by the previous females. It is concluded that the females preferred the parasitized hosts over unparasitized hosts because the benefit of saving time and energy for drilling was more than the cost of progeny survival.     

Intraspecific host discrimination and larval competition inMicroplitis croceipes,Microplitis demolitor,Cotesia kazak (HYM.: Braconidae) andHyposoter didymator HYM: Ichneumonidae), parasitoids ofHeliothis virescens (LEP.: Noctuidae)

Intraspecific host discrimination and larval competition were studied forMicroplitis croceipes (Cresson),Microplitis demolitor Wilkinson,Cotesia kazak (Telenga), andHyposoter didymator (Thunberg), solitary endoparasitoids of the tobacco budworm,Heliothis virescens (F.). In ovipositional choice tests between unparasitized and parasitized hosts, the mean number of ovipositions for unparasitized hosts was significantly higher than the mean number of ovipositions for hosts parasitized once by a conspecific female forC. kazak andH. didymator, demonstrating that females of these two species discriminate against hosts recently (within a few seconds) parasitized by a conspecific female. No significant difference in oviposition occurred between these two kinds of hosts forM. croceipes andM. demolitor. Mean percent parasitization by a second conspecific female was determined at 24, 48, and 72 h delays in time between the first and second female attack, and with no delay. Except for the 0 h time delay forC. kazak andH. didymator, percent parasitization by a second conspecific female generally decreased as the delay in time between the first and second female attack increased. When the second parasitization immediately followed the first, one parasitoid larva always eliminated the other by physical combat. With a 24 or 48 h delay between the first and second parasitization, the younger larva was the victor over the older larva forM. croceipes, M. demolitor andC. kazak in at least 50% of the cases. Elimination of older larvae by younger larva was by physical attack. However, forH. didymator, the older instar was the victor, and elimination of younger larvae by older larvae was probably through physiological processes. Further, older larvae ofH. didymator apparently killed the eggs of the second female by physiological processes.      

Two tachinid species discriminate between parasitized and non-parasitized hosts

The abilities ofMyiopharus doryphorae (Riley) andM. aberrans (Townsend) (Diptera: Tachinidae) to discriminate between parasitized and non-parasitized Colorado potato beetle (host)Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) larvae, were investigated under laboratory and field conditions. Laboratory experiments showed that bothMyiopharus species have a significantly greater frequency for larvipositing in non-parasitized hosts over parasitized ones. Direct field observations of larvipositional behavior of bothMyiopharus species over three growing seasons showed effective restraint from larviposition into parasitized hosts, while larviposition into nonparasitized ones occurred readily. Avoidance of previously-parasitized hosts occurred after the larvipositing flies briefly landed on host larvae without attempting to insert the larvipositor. The low levels of superparasitism which occurred in the caged experiments and in the field appeared to be due to a breakdown of the larvipositing parasitoids' restraint when they met only parasitized hosts or when many parasitoids competed for reduced numbers of hosts late in the season.     

Host Age Response in the Parasitoid Wasp Spalangia cameroni (Hymenoptera: Pteromalidae)

Female Spalangia cameroni produced more offspring from younger house fly pupae, both when given a choice of host ages and when not given a choice. Host age did not affect offspring survivorship. Offspring were larger when they had developed on younger hosts and the effect was independent of offspring sex. Having previously parasitized old hosts versus young hosts did not reduce a female's production of offspring in subsequent hosts. Females distinguished between young and old hosts both in the light and in the dark. Females did not distinguish between host ages prior to physical contact with the host but could distinguish by the time they first began exploring a host by tapping it with their antennae; thus, they could distinguish before drilling into a host.     

Parasitism by Pteromalus cerealellae (Hymenoptera: Pteromalidae) on the Cowpea weevil, Callosbruchus maculatus (Coleoptera: Bruchidae): Host density, temperature effects, and host finding ability

Studies on interactions between a larval parasitoid, Pteromalus cerealellae (Boucek) and one of its hosts, Callosobruchus maculatus (F.) were carried out in the laboratory. The number of host larvae parasitized by P. cerealellae increased with host larvae at low densities and tended to a plateau at a density of 25 larvae per female parasitoid. Each parasitoid was able to parasitize more hosts and produced more offspring at 20 and 25 °C than at 30 °C. The number of non-infested seeds mixed with seeds infested with the last instar of C. maculatus did not preclude P. cerealellae from identifying infested seeds and attacking larvae inside them. When infested seeds were tightly packed, several host larvae escaped parasitism. P. cerealellae may be a useful biological control agent in newly harvested cowpea with low C. maculatus infestation, and lowering the temperature of the storage system may enhance the effectiveness of this parasitoid.     

Recognition of heterospecific parasitism: Competition between Aphidiid (Aphidius ervi) and Aphelinid (Aphelinus asychis) parasitoids of Aphids (Hymenoptera: Aphidiidae; Aphelinidae)

The solitary parasitoids Aphidius erviHaliday (Hymenoptera: Aphidiidae) and Aphelinus asychisWalker (Hymenoptera: Aphelinidae) attacked but generally did not oviposit in pea aphids parasitized by the other species. Wasps selectively oviposited in unparasitized hosts when given a choice. Host discrimination depended on the recognition of internal cues. Females of A. asychiseither could not recognize or ignored A. ervi'sexternal host marking pheromone. Under most conditions, A. ervisurvived in superparasitized hosts, killing competing A. asychislarvae by physical attack and possibly physiological suppression. The outcome of larval competition was not affected by oviposition sequence or age difference between larvae; A. asychissurvived only when it had substantially completed larval development before the host was superparasitized by A. ervi.It is suggested that competition for host resources incurs a cost, for the winner in terms of reduced size or increased development time and for the loser in terms of lost progeny and searching time. Consequently, heterospecific host discrimination can be functional. Internal, and probably general, cues enable wasps to recognize and avoid oviposition in hosts already parasitized by an unrelated species.