Comparison of oviposition behavior of a solitary and a gregarious parasitoid (Hymenoptera: Mymaridae)

Anaphes victus Huber andAnaphes listronoti Huber (Hymenoptera: Mymaridae) are respectively solitary and gregarious egg parasitoids of the carrot weevil,Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae). We made detailed ethograms of the oviposition behavior on unparasitized and parasitized hosts for the two species. We then compared the behavior of virgin and mated females for the oviposition of male and female progenies. The two species did not always oviposit after insertion of the ovipositor, but these punctures without oviposition could be readily differentiated from oviposition.A. victus oviposited only once by puncture, whileA. listronoti deposited one to three eggs during the same sequence. The variability of the duration of the various components was generally lower for a given female than between females. Two components, the abdominal vibrations and the pause, were significantly shorter in ovipositions that resulted in male progency for the two species. However, an important overlap in duration prevents using these differences to sex the progeny at oviposition. Virgin females of both species, although capable of producing only males, exhibited both behaviors. Parasitized hosts were recognized through internal and external markings that were used in host discrimination.     

Responses of a generalist and a specialist parasitoid (Hymenoptera: Eucoilidae) to Drosophilid larval kairomones

Foraging parasitoids are thought to need more specific information than generalists on the presence, identity, availability, and suitability of their insect host species. In the present paper, we compare responses to host kairomones by two phylogenetically related parasitoid species that attack Drosophilidae and that differ in the width of their host range. As predicted, the behavioral response of the parasitoids to host kairomones reflected their difference in host range. The response of the specialist parasitoid Leptopilina boulardiwas restricted to contact kairomones from its natural hosts and one closely related species. In contrast, the generalist parasitoid Leptopilina heterotomaresponded to contact kairomones of a variety of Drosophilidae species.     

The role of host species, age and defensive behaviour on ovipositional decisions in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

The main objective of this study was to determine the extent to which host acceptance behaviour as related to host species, age, and defensive behaviour might explain the differences in host use that exist between two congeneric and sympatric species of parasitic wasps. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). Cotesia species differed in their responses to host species (P. brassicae (L.), P. napi (L.) and P. rapae) and developmental stage (early and late 1st, 2nd and 3rd instars). In no-choice tests, host acceptance by C. rubecula was higher for p. rapae and females did not distinguish among the 6 host ages. In contrast, when foraging for P. brassicae and P. napi, C. rubecula females more readily attacked early first instar. Cotesia glomerata showed a higher degree of behavioural plasticity towards acceptance of Pieris host species and host age than did C. rubecula. Cotesia glomerata females parasitized the three Pieris species and showed higher acceptance of first and second instars over third instar. Oviposition success was also influenced by host defensive behaviour. The frequency and the effectiveness of defensive behaviour rose with increasing age of the host, P. brassicae being the most aggressive Pieris species. Furthermore, the mean duration of C. glomerata oviposition was significantly reduced by the defensive reactions of P. brassicae, which would likely affect parasitoid fitness as oviposition time is positively correlated to clutch size in C. glomerata. Acceptance frequencies corresponded well to field reports of Pieris-Cotesia associations and to patterns of parasitoid larval performance, suggesting that the acceptance phase might be used as a reliable indicator of Cotesia host-specificity.     

The influence of plant species on attraction and host acceptance inCotesia glomerata (Hymenoptera: Braconidae)

Females of the larval parasitoidCotesia glomerata (L.) use plant-associated cues to locate their lepidopteran host,Pieris rapae L. In this study we investigated the influence of four host plant species,Brassica oleracea var.acephala (‘Vates’ kale),Tropaeolum majus (nasturtium),Lunaria annua (honesty), andCleome spinosa (spider flower), on two components of the host selection process inC. glomerata, namely, attraction and host acceptance. Choice tests in a flight tunnel showed that parasitoids were attracted to some host plant species more than to others in the absence of host larvae.B. oleracea was the most attractive plant species, followed byL. annua, T. majus, andC. spinosa. In previous studies it was shown thatB. oleracea carries highly suitable hosts forC. glomerata and that, in the field, parasitization rates on this plant were the highest. When host larvae were reared on the four host plant species and then transferred to a common substrate (B. oleracea var.capitata, cabbage), plant species that had served as diet for the hosts did not have a significant effect on acceptance for parasitization. Thus, parasitoids were attracted to host plant species differentially, but they did not discriminate among host larvae based on the dietary history of their hosts. ForC. glomerata, it appears that phytochemistry mediates host selection more by influencing parasitoid attraction than it does by affecting host acceptance.     

Comparison of learning in related generalist and specialist eucoilid parasitoids

Effects of learning in two microhabitat specialists, Leptopilina boulardia Barbotin et al. and L. fimbriata Kieffer were compared to previous and new results of learning in the microhabitat generalist L. heterotoma Thomson. Females were given one or more oviposition experiences on hosts in different types of substrate. In all species oviposition experience affected the choice for a substrate, although this effect of learning was considerably less in L. fimbriata compared to the other two species. Patch times, known to be highly determined by experience in the generalist L. heterotoma, were much less flexible in the specialists. L. boulardi and L. fimbriata have fixed patch times on their natural substrate and have variable patch times on other substrates only. In all three species one oviposition affected the choice for a substrate. Additional ovipositions showed no different effect. An accumulative effect of the number of ovipositions on patch times was found in L. heterotoma only. Retention of the learning effect was only studied in L. boulardi, and was shown to be similar to that reported for L. heterotoma, i.e. two to three days. Although learning was found in both the generalist and the specialist species studied, it seems to affect their foraging behaviour differently.     

Effects of postemergence experience on searching and landing responses of the insect parasitoid,Cotesia congregata (Say) (Hymenoptera: Braconidae), to plants

Postemergence experience with one of six plant species, in the presence of the host larva, modified the searching response of reproductively mature females of Cotesia congregata(Say) to these plants in at least one of three ways: (1) an increased response to the plant experienced at emergence, (2) an increased response to other plants, or (3) an inhibited response to other plants. Landing and searching responses were differentially affected by postemergence experience. For example, postemergence experience with tobacco (a common plant) in the presence of the host larva induced a landing preference for this plant over parsley (a novel plant) but did not affect searching responses to either plant, whereas experience with parsley and the host larva induced an increased searching response to parsley but a landing preference for tobacco. Differential effects of postemergence experience may reflect the type of stimuli involved in searching or landing and may have adaptive significance.     

Resource Partitioning among Parasitoids (Hymenoptera: Braconidae) of Phoracantha semipunctata in Their Native Range

The Eucalyptus longhorned borer, Phoracantha semipunctata (F.), is native to Australia, but it has been introduced without its natural enemies into many parts of the world in which its Eucalyptus spp. host has been planted. The beetle has developed large populations in these novel habitats and has been responsible for the mortality of large numbers of trees. Although there is a considerable catalogue of the parasitoids of the beetle in Australia, limited ecological information on the assemblage of parasitoids attacking P. semipunctata is available. We removed bark from 40 felled trees, recorded gallery width and bark thickness over parasitized larvae, and removed all parasitoids. Adult size, sex, and species were recorded when the parasitoid pupae eclosed. Syngaster lepidus Brullè, Jarra phoracantha Austin, Quicke, and Marsh, J. maculipennis Austin, Quicke, and Marsh, and J. painei Austin and Dangerfield were most commonly collected. The solitary parasitoid S. lepidus preferred smaller larvae than did the gregarious Jarra spp. The two species with shorter ovipositors, J. maculipennis and J. painei, parasitized larvae under thinner bark than did the other two species with longer ovipositors. There was a significant positive correlation between host larval size and number of parasitoid pupae of the gregarious species. Also, there was a significant positive correlation between host larval size and parasitoid adult size. The ecological relationships between this assemblage of parasitoids and their beetle host may be useful in establishing an effective biological control program.     

Host recognition byPimpla instigator F. (Hymenoptera: Ichneumonidae): Preferences and learned responses

This study identifies some previously unreported tactile and visual cues used by the pupal parasitoid Pimpla instigatorF. (Ichneumonidae) to recognize potential hosts. Paper cylinders were presented to the wasps as simple models of lepidopteran pupae. Acceptance of these models was evaluated by determining the frequency with which the wasps punctured the cylinders with their ovipositors. The length of the cylinders did not influence acceptance of the models; however, both surface texture and structural modifications to the ends of the cylinder did affect the frequency of punctures. Smooth cylinders were punctured more often than roughened cylinders, and cylinders with closed ends were frequently punctured, whereas open-ended cylinders were consistently rejected. The wasps also discriminated between blue and yellow cylinders and could be trained to associate blue or yellow with the presence of hosts. Preferences were established during a single 90- min training period and persisted for at least 4 days following training.     

Factors governing host discrimination byOpius dimidiatus (Ashmead) (Hymenoptera: Braconidae)

The detection of, and response to, parasitized hosts by female parasitoids can involve a number of complex phenomena, including the ability of females to discriminate between unparasitized and parasitized hosts, the physiological state of females, and the nature and longevity of the cues they are detecting. The discriminatory ability of Opius dimidiatus,a solitary endoparasitoid of Liriomyza trifoliiwas examined. Experiments showed, first, that when presented with a choice of unparasitized hosts and hosts which they had previously parasitized, O. dimidiatusfemales could discriminate between them, both upon encounter (before probing) and after probing with the ovipositor. It was further demonstrated that the detection of parasitized hosts was time dependent: females could discriminate on the basis of antennal examination for up to 2.5 h after the first egg was laid. Postprobing discrimination lasted for up to 3.5 h. Some of the causal and evolutionary factors resulting in superparasitism are discussed.     

Chemically mediated associative learning in the host foraging behavior of the aphid parasitoidAphidius ervi (Hymenoptera: Braconidae)

The effect of experience on the responsiveness of the braconid parasitoidAphidius ervi to host(Acyrthosiphon pisum)-associated cues was investigated on bean plants(Vicia faba) using a wind tunnel bioassay. Oviposition experience on the plant-host complex significantly increased the oriented flight and landing responses ofA. ervi females to an undamaged plant and to a plant-host(A. pisum) complex. However, oviposition experience onA. pisum aphids when isolated from the plant did not change their responses to the intact plant and the complex. Searching on an unwashed plant which had been previously damaged byA. pisum also increased their response to an undamaged plant and a host-damaged plant, whereas the experience of searching on an undamaged plant did not significantly change their responses to undamaged plants. However, when parasitoids were allowed to search on an undamaged plant which had been sprayed withA. pisum honeydew, this significantly increased their response to an undamaged plant. Oviposition experience on the plant-host complex and foraging experience on a host-damaged plant or an undamaged plant sprayed with honeydew also significantly reduced the mean time taken by the parasitoids to respond in the wind tunnel. The behavioral changes associated with such experience were acquired within 30 min and persisted for at least 3 days. The results demonstrate the capacity ofA. ervi to learn associatively olfactory cues from plants, host-damaged plants, and plant-host complexes and confirm the role of aphid honeydew as a host recognition kairomone forA. ervi.     

An illustrated key to Neotropical species of the genus Meteorus Haliday (Hymenoptera,Braconidae, Euphorinae)

A comprehensive key for 75 species of Meteorus distributed across 15 Neotropical countries is presented. Eleven new species from Bolivia, Costa Rica and Ecuador are described: Meteorus albistigma, Meteorus carolae, Meteorus eurysaccavorus, Meteorus fallacavus, Meteorus flavistigma, Meteorus haimowitzi, Meteorus magnoculus, Meteorus martinezi, Meteorus microcavus, Meteorus noctuivorus and Meteorus orion. Expanded range distributions are recorded for Meteorus andreae, Meteorus farallonensis, Meteorus guineverae, Meteorus jerodi, Meteorus kraussi, Meteorus papiliovorus and Meteorus quimbayensis. The host of Meteorus jerodi is reported for the first time: a noctuid larva feeding on Asteraceae. Meteorus papiliovorus is recorded attacking Papilionidae larvae in Ecuador, therefore displaying a similar host family preference as formerly documented from Costa Rica and Colombia.     

Host-related fitness trade-offs in a presumed generalist parasitoid, Diaeretiella rapae (Hymenoptera: Aphidiidae)

Abstract.  1. Host ranges of parasitoid wasps are mediated by behavioural responses to hosts and their environment (infectivity), and development in hosts (virulence). Determinants of host range were measured in Diaeretiella rapae (Hymenoptera: Aphidiidae), which has been described as a generalist that attacks more than 60 species.
2. In northern Colorado, this wasp mainly attacks two hosts: cabbage aphid ( Brevicoryne brassicae ) and Russian wheat aphid ( Diuraphis noxia ). Here, laboratory experiments are described in which D. rapae originating from these two hosts were offered several hosts for oviposition. Both infectivity and virulence were measured.
3. Infectivity included host acceptance and handling time, while virulence was measured as productivity (number of progeny), survival of immatures within hosts, development time, and sex ratio.
4. Wasps had higher productivity and survival when attacking 'home' hosts than 'alternate' hosts, and trade-offs were found by quantitative genetic analyses to be genetically determined. Sex ratio and development times also showed trade-offs, but mainly related to the host environment in which females were reared.
5. In previous genetic studies in northern Colorado, populations were genetically subdivided on the scale of 1 km. The fitness differences described here could be strong enough to create populations adapted to different hosts, but it appears that gene flow is sufficient to prevent formation of separate lineages on the two hosts.
6. Rather than being a generalist with a broad host range, D. rapae is a serial specialist, attacking particular hosts according to availability in different seasons or in different geographical areas.     

Role of plant volatiles in host location by the specialist parasitoidMicroplitis croceipes cresson (Braconidae: Hymenoptera)

A study was conducted to determine the primary source of volatile cues within the plant-host complex used by hostseeking freeflying female Microplitis cro-ceipesCresson in flight tunnel bioassays. In single-source and two-choice tests, using wasps given an oviposition experience on either cotton (Gossypium hirsutum)or cowpea (Vigna unguiculata)seedlings damaged by corn earworm (CEW; Helicoverpa zeaBoddie), the damaged seedlings were significantly more attractive than the CEW frass, which was in turn more attractive than the larvae themselves. In a series of two-choice wind-tunnel tests, the discriminatory ability of the wasps was examined, following various oviposition experiences. Significantly more wasps flew to plants with old damage than to plants with fresh damage, regardless of whether they had experience on fresh or old damage. In a comparison of plant species, wasps with only one experience on either hostdamaged cotton or host-damaged cowpea were unable to distinguish between them, and showed no preference for either plant, whereas wasps with multiple experiences on a particular plant preferentially flew to that plant in the choice test. In comparing hosts with nonhosts, wasps successfully learned to distinguish CEW from beet armyworm (BAW; Spodoptera exigua)on cotton but were unable to distinguish CEW from either BAW or cabbage looper (Trichoplusia ni)on cowpea. The results show the important role played by plant volatiles in the location of hosts by M. croceipesand indicate the wasps limitations in discriminating among the various odors. The ecological advantages and disadvantages of this behavior are discussed.     

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.     

中国并腹茧蜂属一新种记述（膜翅目：茧蜂科）

记述采自中国黑龙江的并腹茧蜂属Hylcalosia Fischer 1新种——沟腹并腹茧蜂H.ventisulcata sp.nov..该新种与海氏并腹茧蜂Hylcalosia hymaenei Belokobylskij最为相似,但新种的如下特征容易与后者相区别:1)下颚须相当长,可达后足基节处;2)腹部第2-3节背板具1条宽而浅的中纵沟,几乎纵贯2、3背板;3)前翅2-SR脉基部弯曲,后翅m-cu脉对叉式;4)腹部第1背板中长等于端宽。文中修订了该属全世界已知种的检索表。新种模式标本保存于福建农林大学益虫研究所。     

Postemergence learning in the insect parasitoid,Cotesia congregata (Say) (Hymenoptera: Braconidae)

Exposing newly emerged females of Cotesia congregata(Say) to wild cherry, an inherently unattractive plant, and their host larvae at 0–4 h after adult emergence induced a positive searching response to wild cherry and an inhibited response to cabbage, an attractive plant. Inherent responses were not affected when females were exposed to their hosts at 0–12 h and to cherry at 8–12 h after emergence. The induced response to cherry was constant until its disappearance at 6–7 days;inhibition of the response to cabbage was released at 4–5 days after emergence. Postemergence exposure to cherry and parasitoid cocoons induced similar but weaker searching responses. Induced searching responses exhibit features of associative learning and receptor modification. In addition to its presumed role in foraging, postemergence experience with plants may encourage assortative mating of C. congregatawithin suitable host habitats and, thus, facilitate local adaptations to specific plants.     

Host recognition by the specialist endoparasitoidMicroplitis croceipes (Hymenoptera: Braconidae): Role of host- and plant-related volatiles

The specialist parasitoidMicroplitis croceipes Cresson can parasitize only noctuid larvae in the generaHelicoverpa andHeliothis. To be successful in their search for hosts, the ability to distinguish hosts from nonhosts feeding on the same plant is beneficial. In flight tunnel experiments, we found that prior to landing on the odor sourceM. croceipes were able to distinguish volatiles released from frass of host larvae(Helicoverpa zea Boddie) and nonhost larvae (Spodoptera exigua Hübner andSpodoptera frugiperda J. E. Smith) fed on cotton. However, an initial contact experience with frass of cotton-fed host larvae appeared to be critical for this ability. Wasps that had antennated frass of host larvae fed pinto bean diet were equally attracted to frass of host and nonhost larvae fed on pinto bean diet. In short-range walking experiments, wasps located cotton-fed host larvae faster than diet-fed larvae, regardless of their experience. Wasps that had antennated frass of cotton-fed host larvae were less attracted to cotton-fed nonhost larvae, compared to host larvae, and preferred to sting host larvae. Plant-related volatiles in host frass and larvae appear to play a major role in the successful location of host larvae.     

The parasitoidCotesia glomerata (Hymenoptera: Braconidae) discriminates between first and fifth larval instars of its hostPieris brassicae,on the basis of contact cues from frass,silk, and herbivore-damaged leaf tissue

Adult females of the larval parasitoidCotesia glomerata (L.) respond to chemical cues associated with feeding damage inflicted on cabbage plants by its host,Pieris brassicae (L.). The use of these infochemicals by the parasitoid during selection of the most suitable host instar was investigated. The parasitoid can successfully parasitize first-instar host larvae, while contacts with fifth-instar larvae are very risky since these caterpillars react to parasitization attempts by biting, spitting, and hitting, resulting in a high probability of the parasitoid being seriously injured or killed. Observations of the locomotor behavior of individual wasps on leaves with feeding damage inflicted by the first and the fifth larval instars and on host silk and frass showed that several cues affect the duration of searching by the parasitoids after reaching a leaf: cues on the margin of the feeding damage and cues in the host frass and silk. Whole frass, silk, and hexane extracts of frass obtained from first-instar elicited parasitoid's searching behavior significantly longer than frass, silk, and hexane extract of frass from the fifth instar. The results demonstrate thatC. glomerata can discriminate between first instars, which are more suitable hosts, and fifth instars ofP. brassicae without contacting the caterpillars, by exploiting instar-related cues.     

The role of learning in adult food location by the larval parasitoid,Microplitis croceipes (Hymenoptera: Braconidae)

Wind tunnel experiments were conducted to determine roles of odor learning in food foraging of the larval parasitoid,Microplitis croceipes (Hymenoptera: Braconidae). Females that had neither fed on sucrose water nor experienced any odor and females that had experienced an odor without feeding failed to respond to any odors in a wind tunnel. Most of the females that had fed without an odor also did not respond to odors. However, most of the females that had experienced an odor during feeding on sucrose water flew to the odor. These results indicate that when females experience an odor during feeding, they learn to associate the odor with food and subsequently respond to the odor. As age of females increased, their response to an experienced odor increased, peaked 2 to 5 days after emergence, and then decreased. With an increasing number of odor experiences while feeding, accuracy of females choosing the experienced odor increased. Females that experienced an odor while feeding three to five times chose the experienced odor 90% of the time. When females experienced an odor while feeding five times, the memory of food associated odor lasted at least 2 days. When they experienced food with two odors successively, they could memorize both odors, and multiple experiences did not cause memory interference. Even when females had learned a food-associated odor, their response to the learned odor ceased after several visits on patches containing the odor but no food. Such negative experience may cause switching of food searching to new odors by females.     

Odor learning byLeptopilina boulardi,a specialist parasitoid (Hymenoptera: Eucoilidae)

We studied odor learning in Leptopilina boulardi,a specialist larval parasitoid of Drosophila melanogaster.The behavioral responses of differently experienced females to an artificial odor (Must de Cartier, Paris) were analyzed using a fourarmed airflow olfactometer. The responses of females with an oviposition experience in the presence of the perfume were compared with those of four control groups. As controls we used naive females, females with an oviposition experience in the absence of odor, females which had been previously exposed to perfume but without an oviposition experience, and females with an oviposition experience which also had been exposed to perfume but not at the same time. The results demonstrate that a specialist such as L. boulardican learn very well to respond to an artificial odor by associating this odor with a reward, i.e., an oviposition. The four control groups responded more or less in a similar way.