斑痣悬茧蜂寄生、发育和生殖特征与寄主幼虫日龄的关系

【目的】“圆屋顶形”假说认为,对单寄生性姬蜂和茧蜂适合度而言,中间龄期幼虫寄主的品质高于更早和更晚龄期幼虫。该假说得到许多研究支持,但这些研究常以寄主幼虫脱皮划分虫龄,很少观测生殖特征,从而难以确切和全面描述适合度随寄主生长发育变化而变化的关系。本研究旨在检验“圆屋顶形”假说。【方法】本研究以斜纹夜蛾Spodoptera litura不同日龄幼虫为寄主,观测斑痣悬茧蜂Meteorus pulchricornis寄生和发育特征,并测定成蜂生殖力。【结果】线性回归分析表明,雌蜂对中间日龄寄主幼虫的寄生率大于对两端日龄寄主幼虫的寄生率;蜂卵至成虫的存活、成虫体型大小及其生殖力（产卵量）等适合度相关特征均表现出中间日龄寄主幼虫处理大于两端日龄幼虫处理。【结论】研究结果支持“圆屋顶形”假说。     

Interaction of a solitary larval endoparasitoid, Microplitis mediator, with its host, Mamestra brassicae: host acceptance and host suitability

Abstract:  Microplitis mediator (Haliday) (Hym., Braconidae) is an important parasitoid of early instar larvae of the European cabbage moth, Mamestra brassicae L. (Lep., Noctuidae). In the laboratory, we examined attack responses of female M. mediator to the first three larval instars of M. brassicae . Females were presented with M. brassicae larvae either one individual at a time in a no-choice experiment, or three individuals, one from each instar, simultaneously in a choice experiment. Whether or not there was choice, naïve female parasitoids attacked a high proportion of larvae and did not discriminate among instars. In the no-choice experiment, attacked larvae were reared, and parasitoid cocoons were produced from about 76% of larvae attacked as first and second instars, but from only 19% of larvae attacked as third instars. Dissections of attacked larvae from the choice experiment showed that about 79% of attacks on first and second instars resulted in oviposition compared with only 49% for third instars. When given choice, frequency and number of attacks on first instar larvae increased with increasing parasitoid experience. Our results suggest that first and second instar larvae of M. brassicae are suitable hosts for M. mediator , but that third instar larvae are suboptimal both because oviposition attempts were frequently unsuccessful and because immature parasitoids failed to complete development. Nevertheless, naïve attacking parasitoids exhibited minimal discrimination among instars, although experienced parasitoids most frequently attacked first instar larvae. The host selection behaviour of M. mediator is discussed in the context of optimal foraging theory and implications for biological control.     

Does mother really know best? Oviposition preference reduces reproductive performance in the generalist parasitoid Aphidius ervi

The reproductive success of female parasitoids is dependent on their ability to accurately assess the suitability of a host for larval development. For generalist parasitoids, which utilize a broad range of species and instars as hosts, a set of assessment criteria determines whether a host is accepted or rejected. The suitability of a host, however, can only be imperfectly assessed by the female parasitoid, which can result in the selection of lesser quality hosts for oviposition. In this study we explored the disparity between host quality and host preference using the generalist koinobiotic parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) and the host Aulacorthum solani (Harris) (Homoptera: Aphididae), the foxglove aphid. The second instar hosts produced the highest level of reproductive success, while third and fourth instars resulted in a substantially reduced reproductive performance. When given a choice of host instars, parasitoids preferred the older hosts for oviposition disregarding their reduced suitability for larval development. Results are discussed in context of mechanisms involved in A. ervi host selection and biases in the criteria used to assess hosts that may arise when parasitoids transfer host species between generations.     

Seasonal phenology and stage-specific parasitism of the apple ermine moth, Yponomeuta malinellus Zeller, in Korea

The apple ermine moth, Yponomeuta malinellus Zeller (Lepidoptera: Yponomeutidae), is a tent caterpillar that feeds on Malus spp. in Korea. Populations of the moth in native areas appeared to be regulated by the assemblage of parasitoids. Phenological associations between host stages and parasitoids, susceptible stage(s) of the host for each parasitoid, and stage‐specific parasitism were studied. The egg larval parasitoid Ageniaspis fuscicollis (Dalman) had highest parasitism of first instar larvae (24%), with 14% parasitism of other larval stages. Dolichogenidea delecta (Haliday) was recovered from all larval instars with the highest parasitism rate of second instar larvae (20.1%), followed by 19.9% parasitism of mid‐larval hosts. Herpestomus brunicornis Gravenhorst was reared from second instar larvae through to pupal collection, and had the highest parasitism rate (29.9%) at the pupal stage. The larval pupal parasitoid Zenillia dolosa (Meigen) was recovered from mid‐larval to pupal stages with the highest parasitism rate (5.5%) occurring in third to fourth instar larvae. The host stages for developing A. fuscicollis completely overlap with those of D. delecta, and with those of H. brunicornis to some degree. A statistically significant negative correlation exists between A. fuscicollis and these dominant parasitoids, indicating competitive interaction within the host.     

Diadegma mollipla parasitizing Plutella xylostella: host instar preference and suitability

Oviposition decisions (i.e., host selection and sex allocation) of female parasitoids are expected to correspond with host quality, as their offspring fitness is dependent on the amount and quality of resources provided by a single host. The host size model assumes that host quality is a linear function of host size, with larger hosts believed to contain a greater quantity of resources, and thus be more profitable than smaller hosts. We tested this assertion in the laboratory on a solitary larval–pupal parasitoid Diadegma mollipla (Holmgren) (Hymenoptera: Ichneumonidae) developing on three instars (second–fourth) of one of its hosts, the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae). In a no‐choice test, parasitism levels and sex ratio (i.e., proportion of female progeny) were significantly high in hosts attacked in the second instar followed by third then fourth instars. However, the few parasitoids that completed a generation from the fourth instars did so significantly faster than conspecifics that started development in the other two instars. In direct observations, however, the parasitoids (i) randomly attacked the various host instars, (ii) spent a similar period examining the various host instars with their ovipositors, (iii) subdued all three host instars with about the same effort, and (iv) no statistical differences were observed in the attack rates on the three host instars. In a choice test, the females parasitized significantly more third instars followed by second then fourth instars. However, total parasitism in this experiment was 43% lower compared to parasitism of only second instars in the no‐choice test. No significant differences were detected in progeny sex ratios. In both choice and no‐choice tests, significantly more fourth instars died during the course of the experiments than second instars, while third instars were intermediate. The higher parasitism of third than second instars in the choice test indicates that the females perceived larger hosts as higher quality than smaller hosts, despite their lower suitability for larval development.     

Manipulation of fifth-instar host (Manduca sexta) ecdysteroid levels by the parasitoid wasp Cotesia congregata

Although 5th (last) instar parasitized Manduca sexta larvae undergo developmental arrest and do not wander, they exhibit a small hemolymph ecdysteroid peak (300-400pg/&mgr;l) which begins one day prior to the parasitoid's molt to the 3rd (last) instar and concomitant emergence from the host. Ecdysteroids present in this peak were 20-hydroxyecdysone, 20,26-dihydroxyecdysone and one or more very polar ecdysteroids, as well as small amounts of 26-hydroxyecdysone and ecdysone. In parasitized day-1 and -2 5th instars ligated just behind the 1st abdominal proleg, hemolymph ecdysteroid levels increased in both anterior and posterior portions (100-500pg/&mgr;l), while in unparasitized larvae, hormone levels only increased in the anterior portion (100-350pg/&mgr;l). Thus, the ecdysteroid peak observed in host 5th instars was probably produced, at least in part, by the parasitoids. It may serve to promote Cotesia congregata's molt from the second to the third instar and/or to facilitate parasitoid emergence from the host. In parasitized day-1 and -2 5th instars ligated between the last thoracic and 1st abdominal segments, hemolymph ecdysteroid titers reached much higher levels (500-3500pg/&mgr;l) in the anterior portion (no parasitoids present) than in the posterior portion (150-450pg/&mgr;l). Therefore, it appears that the parasitoid's regulation of hemolymph ecdysteroid titers occurs at two levels. First, parasitization neutralizes the host's ability to maintain its normal hemolymph ecdysteroid levels. Second, in a separate action, the parasitoid manipulates the ecdysteroid-producing machinery so that hemolymph levels are maintained at the 200-400pg/&mgr;l characteristic of day 3-4 hosts. This is the first report of a parasitoid's ability to interfere with the normal inhibitory mechanisms which prevent prothoracic gland production of ecdysteroid at inappropriate periods of insect growth and development.     

Reproductive biology and small-scale rearing of cherry bark tortrix and its candidate biological control agent

Abstract:  The larval parasitoid, Campoplex dubitator (Hym.,: Ichneumonidae), is under consideration as an agent for classical biological control of cherry bark tortrix (CBT), Enarmonia formosana (Lep.,: Tortricidae), in North America. A comprehensive risk analysis of the candidate agent will require prosperous cultures of both the pest and its parasitoid. We present a rearing method for small-scale production of both species using a bean-based artificial diet, with additional information on the reproductive biology of C. dubitator . Based on estimated survivorship probabilities, a CBT egg had a 70% chance of developing completely to the adult stage under this system. The success of parasitism, however, was very dependent on the instar of the CBT host larva at the time of oviposition. All parasitised first instar larvae died shortly after the attack, rendering them unsuitable for oviposition, while 50% of parasitised second instar larvae died prematurely. In contrast, early mortality was 15–30% for larvae parasitised in the third to fifth instars. Regardless of the instar at oviposition, approximately 90% of the surviving hosts yielded parasitoids, showing a high acceptance by C. dubitator of second to fifth instars for oviposition.     

Effects of host instar and size on parasitization efficiency and life history parameters ofOpius dissitus

The braconid parasitoidOpius dissitus Muesebeck (Hymenoptera, Braconidae) produced 1.7 to 3 times more offspring when provided second and third instar leafminers (Liriomyza sativae Blanchard) as compared to first instars. Females arising from parasitization of different instars did not differ significantly in numbers of chorionated eggs in their ovaries at adult eclosion. Development time was prolonged by about two days when parasitoid oviposition occurred in first, as compared to third instar hosts. Parasitoid length was positively correlated with host weight (r²=0.75). Because only 7% of variation in host weight could be explained by host density, parasitoid length varied considerably among hosts reared at the same density. Longevity and lifetime fecundity of parasitoids were inversely related to the weight of their hosts.     

Regulation ofHelicoverpa zea larval behavior by the parasitoidEucelatoria bryani

The parasitoidEucelatoria bryani Sabrosky regulates the larval behavior of its hostHelicoverpa zea (Boddie). Parasitized third, fourth and fifth instars burrow into the soil 0.7–3.4 days earlier than unparasitized larvae that normally enter the soil to pupate at the end of the fifth and final larval instar. Parasitized third instars molt once then burrow as fourth instars, one instar earlier than normal. WhenE. bryani pupariated on the soil surface in the field, none survived to the adult stage. However,E. bryani adults emerged from 49.2% of hosts that had burrowed into the soil. By accelerating the timing ofH. zea burrowing behavior and causing host larvae to enter the soil before death,E. bryani ensures its pupariation in an environment with improved protection against natural enemies and lethal temperatures.     

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.     

Biology of Microplitis kewleyi (Hymenoptera: Braconidae): A parasite of Agrotis ipsilon (Lepidoptera: Noctuidae) larvae

Microplitis kewleyi Muesebeck is a gregarious internal parasite of larvae of the black cutworm Agrotis ipsilon (Hufnagel). Studies of the biology of the parasite revealed that there was an inverse relationship between host instar and parasite preference. Duration of development from egg to pupa ranged from 18 days at 27°C to 68.7 days at 16°C. Development from egg to pupa took 13.5–21.6 days when fourth and first instar host larvae, respectively, were parasitized. A larger number of parasites emerged from hosts parasitized in the fourth instar (22.4) than the first instar (11.5). Parasite pupation occurred when the host was in the fifth/sixth instar, depending on the instar parasitized. Thirty‐nine per cent of host larvae exposed as first instars to parasites died before parasite emergence. This decreased to 0% for host larvae exposed as fourth instars. The sex ratio was 1:1.2 (M:F). Thirty‐seven per cent of hosts exposed diurnally were stung, compared to 24% exposed nocturnally. Mean daily progeny was highest (12) on the first day, decreasing to zero after 20 days. Percent host parasitism was also highest on the first day (35%) decreasing to nearly 0% after 18 days. There appear to be three parasite larval instars. Host larvae often remained alive after parasite emergence.     

Parameters influencing the acquisition of competence for metamorphosis in imaginal disks of Drosophila

When imaginal disks from first and early second instar larvae of Drosophila are transplanted into larval hosts that are ready to pupate, they are unable to differentiate adult structures. The disks gradually become competent to respond with imaginal differentiation towards the end of the second larval instar (Fig. 1). The first sign of imaginal differentiation is a light-orange pigment that appears in the presumptive eye region when eye-antennal disks from early second instar larvae were subjected to immediate metamorphosis. This pigment was identified as being composed of ommochromes and drosopterins.Incompetent eye-antennal disks from early second instar donors were cultured in adult females for 2 to 5 days, and then retransplanted into late third instar larval hosts. If the adult host flies were kept on standard food the disks grew by cell multiplication (Fig. 2c) and became competent to undergo imaginal differentiation (Fig. 3). If, on the other hand, the adult hosts were starved on a protein-free sugar diet, cell divisions were effectively blocked in the disks. These did not noticeably grow (Fig. 2b) and remained incompetent (Fig. 3). The block caused by starvation proved to be reversible. Based on these results the hypothesis is advanced that the acquisition of competence requires a minimum number of cell divisions to take place in the disk primordium.     

Effect of entomopoxvirus infection of the smaller tea tortrix, Adoxophyes sp. on the development of the endoparasitoid, Ascogaster reticulatus

Infection of Adoxophyes sp. (Lepidoptera: Tortricidae) larvae by an entomopoxvirus (AsEPV) adversely affected the development of the endoparasitoid, Ascogaster reticulatus Watanabe (Hymenoptera: Braconidae). Parasitoid larvae developing in AsEPV-infected hosts grew more slowly and spent more time in their hosts than did parasitoid larvae developing in noninfected hosts. Percentages of emergence of larval parasitoids that developed in AsEPV-infected hosts were significantly lower than those of parasitoids that developed in noninfected hosts. Parasitoid larvae in AsEPV-infected host perished when their hosts died of AsEPV infection. Significant numbers of parasitized and infected larvae exhibited apolysis to the final instar, whereas noninfected-parasitized larvae died in the penultimate instars due to emergence of parasitoids.     

Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

斑痣悬茧蜂对斜纹夜蛾不同虫龄幼虫的选择及后代发育表现

为探究容性寄生蜂对不同龄期寄主幼虫的选择性及其子代蜂发育表现的关联,通过双选试验观察了斑痣悬茧蜂(Meteorus pulchricornis)对斜纹夜蛾(Spodoptera litura)不同虫龄幼虫的寄生选择,并观察了与子代蜂适合度相关的特性表现.结果表明:斑痣悬茧蜂在4龄与5龄之间未表现出偏好,在2龄和3龄、3龄和4龄之间显著偏好较低虫龄;结茧率在不同寄主虫龄间无显著差异,羽化率随寄主虫龄增大而减小,寄生2、3龄幼虫的子代蜂显著高于寄生5龄;寄生5龄幼虫的子代蜂死亡率比寄生2龄的高2.5倍,比寄生3龄的高5.4倍.寄生3龄幼虫的子代蜂发育历期最短(11.9 d),比寄生4龄幼虫的短6.8 d,比寄生2龄幼虫的短4.7 d;子代蜂体型大小在寄生的寄主虫龄间无显著差异.根据研究结果推测,斑痣悬茧蜂在寄生时可能不是根据寄主龄期来评价寄主品质,而是基于寄主体型大小进行评价.     

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.     

The effect of larval nutrition on egg production in Rhodnius prolixus

Rhodnius prolixus adults which had fed to repletion at each larval instar were frequently autogenous, while those allowed to take only partial blood meals at the 3rd, 4th and 5th larval instars were always anautogenous.Bugs fed to repletion at each larval instar had large abdominal fat bodies throughout adult life. In such bugs, the quantity of blood associated with the production of one egg was small and constant during the course of several blood meals.Bugs fed partial blood meals as larvae had small abdominal fat bodies following ecdysis to the adult; the quantity of blood associated with the production of one egg was large following the first adult feed because some of the blood meal was used to increase the size of the abdominal fat bodies.     

Parasitoid–Pathogen–Pest Interactions of Chelonus insularis, Campoletis sonorensis, and a Nucleopolyhedrovirus in Spodoptera frugiperda Larvae

In this study we examined interactions between two solitary endoparasitoids, the braconid Chelonus insularis and the ichneumonid Campoletis sonorensis, and a multiple-enveloped nucleopolyhedrovirus infecting Spodoptera frugiperda larvae. We examined whether ovipositing females minimize interference by discriminating amongst hosts and examined the outcome of within-host competition between parasitoid species and between the parasitoids and the virus. The egg–larval parasitoid Ch. insularis did not discriminate between virus-contaminated and uncontaminated S. frugiperda eggs; all S. frugiperda larvae that emerged from surface-contaminated eggs died of viral infection prior to parasitoid emergence. The larval parasitoid C. sonorensis also failed to discriminate between healthy and virus-infected S. frugiperda larvae or between larvae unparasitized or parasitized by Ch. insularis. Host larvae parasitized in the egg stage by Ch. insularis were suitable for the development of C. sonorensis when they were multiparasitized by C. sonorensis as first, second, third, and fourth instars, whereas emergence of Ch. insularis was dramatically reduced (by 85 to 100%) in multiparasitized hosts. Nonspecific host mortality was significantly higher in multiparasitized hosts than in singly parasitized hosts. The development time and sex ratio of C. sonorensis in multiparasitized host larvae were unaffected by the presence of Ch. insularis larval stages. Both Ch. insularis parasitized and nonparasitized larvae of the same instar (second, third, or fourth instars) had a similar quantitative response to a challenge of virus inoculum. All host larvae that ingested a lethal dose of virus were unsuitable for Ch. insularis development. In contrast, C. sonorensis did not survive in hosts that ingested a lethal virus dose immediately after parasitism, but parasitoid survival was possible with a 2-day delay between parasitism and viral infection and the percentage of parasitoid emergence increased significantly as the interval between parasitism and viral infection increased. The development time of C. sonorensis was significantly reduced in virus-infected hosts compared to conspecifics that developed in healthy hosts. C. sonorensis females that oviposited in virus-infected hosts did not transmit the virus to healthy hosts that were parasitized subsequently. Field applications of virus for biocontrol of S. frugiperda may lead to substantial mortality of immature parasitoids, although field experiments have not yet demonstrated such an effect.     

The effects of host age and superparasitism by the parasitoid, Microplitis rufiventris on the cellular and humoral immune response of Spodoptera littoralis larvae

To study the dynamics of stage-dependent immune responses in Spodoptera littoralis (Boisd.) larvae (Lepidoptera: Noctuidae), single and superparasitism experiments were carried out using the parasitoid Microplitis rufiventris Kok. (Braconidae: Hymenoptera). Compared to younger (preferred) host larvae, the older (non-preferred) host larvae displayed a vigorous humoral response that often damaged and destroyed the single wasp egg or larva. Superparasitism and host age altered both the cellular and humoral immune responses. Younger host larvae showed a stronger encapsulation response compared to older host larvae. Moreover encapsulation rates in younger hosts (e.g., second instar) decreased with increasing numbers of parasitoid eggs deposited/larvae. In older larvae, the encapsulation rate was low in fourth, less in fifth and absent in sixth instar hosts. Conversely, the order and magnitude of the cellular immune response in S. littoralis hosts were highest in second instar larvae with the first instar larvae being a little lower. The immune response steadily decreased from the third through to the fifth instar and was least obvious in the sixth instar. In contrast, the general humoral immune response was most pronounced in sixth instar larvae and diminished towards younger stages. The results suggest that both cellular and humoral responses are stage-dependent. Wasp offspring in younger superparasitized host larvae fought for host supremacy with only one wasp surviving, while supernumerary wasp larvae generally survived in older superparasitized larvae, but were unable to complete development. Older instars seem to have a method for immobilizing/killing wasp larvae that is not operating in the younger instars.