The effects of host weight at parasitism on fitness correlates of the gregarious koinobiont parasitoid Microplitis tristis and consequences for food consumption by its host, Hadena bicruris

Gregarious koinobiont parasitoids attacking a range of host sizes have evolved several mechanisms to adapt to variable host resources, including the regulation of host growth, flexibility in larval development rate, and adjustment of clutch size. We investigated whether the first two mechanisms are involved in responses of the specialist gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) to differences in the larval weight and parasitoid load of its host Hadena bicruris Hufn. (Lepidoptera: Noctuidae). In addition, we examined the effects of parasitism on food consumption by the host. Parasitoids were offered caterpillars of different weight from all five instars, and parasitoid fitness correlates, including survival, development time, and cocoon weight, were recorded. Furthermore, several host growth parameters and food consumption of parasitized and unparasitized hosts were measured. Our results show that M. tristis responds to different host weights by regulating host growth and by adjusting larval development rate. In hosts with small weights, development time was increased, but the increase was insufficient to prevent a reduction in cocoon weight, and as a result parasitoids experienced a lower chance of successful eclosion. Cocoon weight was negatively affected by parasitoid load, even though host growth was positively affected by parasitoid load, especially in hosts with small weights. Later instars were more optimal for growth and development of M. tristis than early instars, which might reflect an adaptation to the life‐history of the host, whose early instars are usually concealed and inaccessible for parasitism on its food plant, Silene latifolia Krause (Caryophyllaceae). Parasitism by M. tristis greatly reduced total host food consumption for all instar stages. Whether plants can benefit directly from the attraction of gregarious koinobiont parasitoids of their herbivores is a subject of current debate. Our results indicate that, in this system, the attraction of a gregarious koinobiont parasitoid can directly benefit the plant by reducing the number of seeds destroyed by the herbivore.     

Host ontogeny determines parasitoid use of a forest caterpillar

For most organisms, patterns of natural enemy‐mediated mortality change over the course of development. Shifts in enemy pressure are particularly relevant for organisms that exhibit exponential growth during development, such as juvenile insects that increase their mass by several orders of magnitude. As one of the dominant groups of insect herbivores in most terrestrial plant communities, larval lepidopterans (caterpillars) are host to a diverse array of parasitoids. Previous research has described how the frequency of herbivore parasitism varies among host plants or habitats, but much less is known about how parasitism pressure changes during host development. To test whether the two major parasitoid taxa, wasps and flies, differentially attack shared hosts based on host developmental stage, we simultaneously exposed early‐ and late‐instar Euclea delphinii Boisduval (Lepidoptera: Limacodidae) caterpillars to parasitism in the field. We found strong evidence that parasitoids partition hosts by size; adult female wasps preferentially parasitized small caterpillars, whereas adult female flies preferred to attack large caterpillars. Our results demonstrate that host ontogeny is a major determinant of parasitoid host selection. Documenting how shifts in enemy pressure vary with development is important to understanding both the population biology and evolutionary ecology of prey species and their enemies.     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.     

Rearing temperature and parasitoid load determine host and parasitoid performance in Manduca sexta and Cotesia congregata

1. Temperature strongly influences the rates of physiological processes in insects, including the herbivore Manduca sexta and its larval endoparasitoid Cotesia congregata. Parasitisation by C. congregata decreases the growth and consumption of food by larval M. sexta. However, the effects of temperature on parasitised caterpillars and the developing wasp larvae are largely unknown. 2. In this study, parasitised and unparasitised caterpillars were reared at three constant temperatures (20, 25 and 30 °C) throughout larval development. Caterpillar mass gain and consumption were monitored daily until wandering (unparasitised control group) or wasp emergence (parasitised group) was observed. Development time and survival to emergence were measured as metrics of parasitoid performance. 3. Parasitised M. sexta developed more slowly than unparasitised controls, but had similar cumulative consumption until the terminal instar. Parasitised caterpillars with relatively large parasitoid loads had higher rates of consumption and growth than those with smaller loads. Both temperature and parasitoid load strongly affected wasp success. Mean development time to wasp emergence increased with low temperatures and with large loads. The combination of warm temperature and large parasitoid loads greatly reduced wasp survival. 4. These results demonstrate the interactive effects of rearing temperature and parasitisation on host consumption and growth rates throughout larval development. In addition, wasp performance was affected by the interaction of temperature and parasitoid load size. High temperatures alter the dynamics of the interaction between the parasitoid and its caterpillar host, which could have far-reaching impacts as the global temperatures continue to rise.     

Growth,development, and behaviour of the parasitised and unparasitised larvae of a shelter‐building moth and consequences for the resulting koinobiont parasitoid

Most attention to size‐time trade‐offs of insects has focused on herbivore risk, with considerably less attention paid to parasitoids. Here, we focus on parasitoid risk, comparing the fates of unparasitised herbivore hosts and parasitised hosts that protect the parasitoids. Success of a koinobiont parasitoid (host grows after parasitisation) depends on maintaining a delicate balance with its host, thereby ensuring its own survival while the host grows. To evaluate growth rate–mortality rate relationships of host and parasitoid, we compared several aspects of the growth, phenology, and behaviour of unparasitised fern moth [Herpetogramma theseusalis (Walker) (Lepidoptera: Crambidae)] larvae and larvae parasitised by Alabagrus texanus (Cresson) (Hymenoptera: Braconidae), a solitary koinobiont (one parasitoid per host) wasp. Host larvae feed and construct shelters on sensitive fern, Onoclea sensibilis L. (Dryopteridaceae). Alabagrus texanus parasitise early‐instar moths in late summer, which overwinter in their host, emerging in mid‐summer to pupate and eclose. During the autumn following hatching and the immediately following spring, parasitised and unparasitised moth larvae did not differ in size, took similar time to choose between satisfactory and unsatisfactory foods, and built similar shelters. Prior to any other changes noted, more parasitised than unparasitised larvae also died when severely starved. Parasitised larvae subsequently grew less and pupated later than unparasitised ones (small size, slow growth), but consumed similar amounts of food. Although the numerically dominant parasitoid of fern moths, we concluded that A. texanus do not efficiently exploit their hosts.     

Cotesia plutellae parasitizing Plutella xylostella: Host-age dependent parasitism and its effect on host development and food consumption

The braconid Cotesia plutellae(Kurdjumov) (Hymenoptera: Braconidae) is amajor solitary, larval endoparasitoid of thediamondback moth, Plutella xylostella(L.) (Lepidoptera: Plutellidae). Parasitism oflarvae of different host instars and fourdevelopmental ages of the 4th instar ofthe pest was examined. The effects of hostinstar at initial parasitization on thedevelopment, survival, size and fecundity ofthe parasitoid were determined in thelaboratory at 25 °C. The effects ofparasitism on host development and foodconsumption were investigated at 28 °C.Cotesia plutellae could parasitize larvaeof all four instars of P. xylostella, butpreferred 2nd and 3rd instars. In achoice test, the relative parasitism indicesfor 2nd, 3rd and 4th instarswere 0.37, 0.39 and 0.24, respectively.Parasitism decreased sharply with increasinghost age in the 4th instar and approachedzero in host larvae that had gone beyond 37%of 4th stadium. The development time andthe final adult size of the parasitoid variedwith the host instar at initial parasitization.Parasitoids with initial parasitism in the4th instar hosts had the shortestdevelopment time, followed by those in the3rd instar, and then by those in the2nd instar. Parasitoids startingparasitism in 2nd instar hosts weresmaller in body size than those starting in the3rd or 4th instar. However, resultantfemales starting parasitism in 3rd instarhosts had the highest fecundity. Parasitizedlarvae exhibited longer development time andincreased food consumption compared withunparasitized ones. This study presents thefirst record that a solitary parasitoidregulates host behavior leading to an increasein food consumption by the host.     

Development of the parasitoid,Cotesia rubecula (Hymenoptera: Braconidae) in Pieris rapae and Pieris brassicae (Lepidoptera: Pieridae): evidence for host regulation

Several recent models examining the developmental strategies of parasitoids attacking hosts which continue feeding and growing after parasitism (=koinobiont parasitoids) assume that host quality is a non-linear function of host size at oviposition. We tested this assumption by comparing the growth and development of males of the solitary koinobiont endoparasitoid, Cotesia rubecula, in first (L1) to third (L3) larval instars of its preferred host, Pieris rapae and in a less preferred host, Pieris brassicae. Beginning 3 days after parasitism, hosts were dissected daily, and both host and parasitoid dry mass was determined. Using data on parasitoid dry mass, we measured the mean relative growth rate of C. rubecula, and compared the trajectories of larval growth of the parasitoid during the larval and pupal stages using non-linear equations. Parasitoids generally survived better, completed development faster, and grew larger in earlier than in later instars of both host species, and adult wasps emerging from P. rapae were significantly larger than wasps emerging from all corresponding instars of P. brassicae. During their early larval stages, parasitoids grew most slowly in L1 P. rapae, whereas in all other host classes of both host species growth to pupation proceeded fairly uniformly. The growth of both host species was markedly reduced after parasitism compared with controls, with the development of P. brassicae arrested at an earlier stage, and at a smaller body mass, than P. rapae. Our results suggest that C. rubecula regulates certain biochemical processes more effectively in P. rapae than in P. brassicae, in accordance with its own nutritional and physiological requirements. Furthermore, we propose that, for parasitoids such as C. rubecula, which do not consume all host tissues prior to pupation, that parasitoid size and host quality may vary independently of host size at oviposition and at larval parasitoid egression.     

Development of a solitary koinobiont hyperparasitoid in different instars of its primary and secondary hosts

Parasitoid wasps are excellent organisms for studying the allocation of host resources to different fitness functions such as adult body mass and development time. Koinobiont parasitoids attack hosts that continue feeding and growing during parasitism, whereas idiobiont parasitoids attack non-growing host stages or paralyzed hosts. Many adult female koinobionts attack a broad range of host stages and are therefore faced with a different set of dynamic challenges compared with idiobionts, where host resources are largely static. Thus far studies on solitary koinobionts have been almost exclusively based on primary parasitoids, yet it is known that many of these are in turn attacked by both koinobiont and idiobiont hyperparasitoids. Here we compare parasitism and development of a primary koinobiont hyperparasitoid, Mesochorus gemellus (Hymenoptera: Ichneumonidae) in larvae of the gregarious primary koinobiont parasitoid, Cotesia glomerata (Hymenoptera: Braconidae) developing in the secondary herbivore host, Pieris brassicae (Lepidoptera: Pieridae). As far as we know this is the first study to examine development of a solitary primary hyperparasitoid in different stages of its secondary herbivore host. Pieris brassicae caterpillars were parasitized as L1 by C. glomerata and then these parasitized caterpillars were presented in separate cohorts to M. gemellus as L3, L4 or L5 instar P. brassicae. Different instars of the secondary hosts were used as proxies for different developmental stages of the primary host, C. glomerata. Larvae of C. glomerata in L5 P. brassicae were significantly longer than those in L3 and L4 caterpillars. Irrespective of secondary host instar, every parasitoid cluster was hyperparasitized by M. gemellus but all only produced male progeny. Male development time decreased with host stage attacked, whereas adult male body mass did not, which shows that M. gemellus is able to optimally exploit older host larvae in terms of adult size despite their decreasing mass during the pupal stage. Across a range of cocoon masses, hyperparasitoid adult male body mass was approximately 84% as large as primary parasitoids, revealing that M. gemellus is almost as efficient at exploiting host resources as secondary (pupal) hyperparasitoids.     

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.     

Factors affecting growth in the koinobiont endoparasitoid Venturia canescens in the flour moth Ephestia kuehniella

With resistance of insect pests to synthetic pesticides on the increase, the role of parasitoid wasps as biological control agents is expanding in pest and resistance management strategies. One of the predictors of reproductive success of endoparasitoids is the relative size of the wasp at host emergence. While in idiobiont parasitoids, where the host stops feeding after parasitism, the wasp size is determined by the host size at the time of parasitism; the size of koinobiont wasps, where the host continues to feed after parasitism, is dependent on additional factors. Here we show that the host mass and temperature are important factors that determine survival and development of the koinobiont endoparasitoid Venturia canescens in late instar larvae of the flour moth Ephestia kuehniella.     

Recruitment of native parasitoids by an exotic leaf miner, Cameraria ohridella: host-parasitoid synchronization and influence of the environment

Abstract

• 1 The horse‐chestnut leaf miner, Cameraria ohridella, is a moth of unknown origin that has recently invaded Europe and severely defoliates the European horse‐chestnut, an important ornamental tree.
• 2 Several indigenous parasitoids have colonized this new host, but parasitism remains low. One of the hypotheses suggested to explain the low parasitism is that candidate parasitoids emerge too early in spring to attack the first host generation and, thus, need early‐occurring leaf miners as alternate hosts. This hypothesis was tested by observing the synchronization between the phenology of the moth and that of its main parasitoids, and by comparing parasitism rates and parasitoid richness in different environments with various levels of biological diversity.
• 3 In spring, the bulk of the parasitoids emerge at least 5 weeks before the occurrence of the first suitable larvae of C. ohridella whereas most parasitoid adults reared outdoors die within 5 weeks after emergence.
• 4 Parasitism rates and parasitoid richness do not increase with biological diversity, suggesting that most parasitoids attacking the first generation of C. ohridella do not come from alternate hosts. Parasitism does not increase later in the year in the subsequent generations, when host‐parasitoid synchronization becomes less critical.
• 5 We conclude that, although the spring emergence of parasitoids is not synchronized with the phenology of C. ohridella, the parasitoids attacking the first generation are probably old or late‐emerging adults of the overwintering generation. The lack of synchronization is probably not the only reason for the poor recruitment of native parasitoids by C. ohridella.

     

Predicting the impacts of climate change on the biological control of Plutella xylostella by Diadegma semiclausum

1. The parasitoid Diadegma semiclausum Hellén (Hymenoptera: Ichneumonidae) has been widely adopted as a biological control agent for Plutella xylostella L. (Lepidoptera: Plutellidae) over the last 80 years. Earlier studies have found differential responses to temperature between the host and its parasitoid and demonstrated the multiple ways in which the parasitoid is more susceptible to elevated temperatures.
2. Using data from experimental studies, the modelling package CLIMEX was used to investigate the suitability of current climates for the host and its parasitoid and the effects on their potential global geographical distributions. The study was then extended to investigate possible changes to these distributions that might result under different climate change scenarios by 2080. The models predict that the global distributions of both the host and parasitoid will be reduced. These changes will not be proportionate and many areas in tropical, sub-tropical, and temperate regions that are currently suitable for D. semiclausum are predicted to become unsuitable for the parasitoid, whereas retaining suitability for P. xylostella. The seasonal dynamics of both the host and parasitoid are also predicted to be significantly reshaped under climate change.
3. Analysis of associations between annual P. xylostella outbreaks and weather conditions in three provinces in China with field data collected between 1995 and 2017 indicated significant effects of temperature on P. xylostella outbreaks at the beginning of the peak season in warmer provinces where P. xylostella can overwinter; such associations were not found in the colder provinces where it is unable to survive harsh winters.

     

Parasitism and constitutive defence costs to host life‐history traits in a parasitoid–host interaction

1. The level of an organism's investment in defences against natural enemies depends on the fitness costs of resisting parasitism and on the costs of maintaining defences in the absence of infection. Heritable variation in resistance suggests that costs exist, but very little is known about the nature or magnitude of these costs in natural populations of animals. 2. A powerful technique for identifying trade‐offs between fitness components is the study of correlated responses to artificial selection. We selected for increased resistance in the Indian meal moth, Plodia interpunctella, following parasitism by the koinobiont parasitoid, Venturia canescens, and measured the cost of resistance to parasitism and the cost of maintaining resistance in the absence of immune challenge during the next generation. 3. Parasitism decreased larval host size, growth, and developmental time and was significantly negatively correlated with the size of surviving host adults. Larvae of the next generation also had a reduced developmental period, whilst the duration of the invulnerable pupal instar was increased. There was no effect on host adult size and related fecundity in the F₁ generation.     

Host stage selection of the mealybug parasitoid Anagyrus spec. nov near sinope

The mealybug parasitoid Anagyrus spec. nov near sinope (Hymenoptera: Encyrtidae) is an undescribed parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Homoptera: Pseudococcidae). We investigated the preference of Anagyrus spec. nov near sinope for six developmental stadia (first‐ and second‐instar nymphs, third‐instar immature females, third‐ or fourth‐instar immature males, pre‐reproductive adult females, and ovipositing adult females) of P. madeirensis and the fitness consequences of the host stage selection behavior. In the no‐choice test, Anagyrus spec. nov near sinope parasitized and completed development in all host stadia except third‐instar immature males. When all host stadia were offered simultaneously, the parasitoids preferred third‐instar immature and pre‐reproductive adult females. Dissection of the stung mealybugs revealed that the clutch size (number of eggs per host) was approximately four and three in the third‐instar and pre‐reproductive females, respectively, and one egg per first‐instar nymph. Parasitoids emerged from P. madeirensis parasitized at third‐instar or pre‐reproductive adult female completed development in the shortest duration, achieved a higher progeny survival rate, larger brood and body size, and the lowest proportion of males. We showed that the continued development of mealybugs had significant influence on the fitness of the parasitoids. Although deposited as eggs in first‐ or second‐instar nymphs, parasitoids emerged from mummies that had attained third‐instar or adult development achieved similar progeny survival rate, brood size, body size, and sex ratio as those parasitoids deposited and developed in third‐instar or adult mealybugs. By delaying larval development in young mealybugs, Anagyrus spec. nov near sinope achieved higher fitness by allowing the parasitized mealybugs to grow and accumulate body size and resources. We suggest that the fitness consequence of host stage selection of a koinobiont parasitoid should be evaluated on both the time of parasitism and the time of mummification.     

Growth rate adjustment of two Drosophila parasitoids in response to the developmental stage of hosts

1. Generalist koinobiont parasitoids often exhibit high flexibility in their development; their larvae shorten or prolong the developmental period depending on the host quality at parasitisation. However, flexibility of the growth rate of parasitoid larvae has rarely been investigated so far. 2. This study investigated how the koinobiont parasitoid wasps Asobara japonica and Leptopilina ryukyuensis regulate their larval growth when they parasitise host Drosophila larvae with varying larval periods. 3. In both parasitoid species, the preimaginal period was longer when they parasitised 1‐day‐old larvae of Drosophila rufa than when they parasitised older larvae of D. rufa or when they parasitised larvae of Drosophila simulans, a species with a shorter larval period than D. rufa. After host pupariation, A. japonica accelerated its growth, thereby showing a biphasic growth curve. On the other hand, L. ryukyuensis did not accelerate its growth after host pupariation. 4. Growth retardation of parasitoid larvae in 1‐day‐old D. rufa larvae would contribute to avoiding excess growth before host pupariation, because the excess growth of parasitoid larvae would have negative effects on host growth. The growth rate acceleration of A. japonica after host pupariation suggests that they enhance resource utilisation in a host that has reached maximum body mass. It remains uncertain as to why L. ryukuensis does not show clear accelerated growth after host pupariation. Nonetheless, these results suggest that parasitoid larvae have the ability to detect the developmental stage of hosts in a species‐specific manner.     

Host plants of the herbivorous insect Mythimna separata (Lepidoptera: Noctuidae) affect its susceptibility to parasitism by the larval parasitoid Cotesia kariyai (Hymenoptera: Braconidae)

Previous studies have shown that the food plant species consumed by herbivorous insects affect their parasitism by parasitoid wasps or flies, but this phenomenon has only been observed in a limited number of systems. Here, we investigate how feeding on different plant species affects the survival of the rice armyworm Mythimna separata (Walker) (Lepidoptera: Noctuidae) and its successful parasitism by the larval endoparasitoid Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae). Survival rate and pupal weight of unparasitised M. separata fed with daikon Raphanus sativus L. var. longipinnatus was lower than with maize Zea mays L. In addition, caterpillars fed with daikon were less susceptible to parasitism by C. kariyai than those fed with maize, and parasitised M. separata larvae fed with daikon had higher survival rates than those fed with maize. These results indicate that while daikon is not an optimal food for unparasitised caterpillars, it may protect the larvae from death by C. kariyai parasitism.     

Flexible larval development and the timing of destructive feeding by a solitary endoparasitoid: an optimal foraging problem in evolutionary perspective

1. In studying the evolution of life-history strategies in parasitoids, considerable attention has been paid to the relationship between host quality and parasitoid fitness. Various workers have reported that host quality influences parasitoid size, development time, and survival. Because body size is frequently correlated with fecundity, longevity, and host-finding ability in parasitoids, this parameter is usually considered to be the main target of selection. 2. In koinobiont parasitoids that consume the entire host before pupation, adult parasitoid size and development time are often strongly correlated with host size at the time when it is developmentally arrested through destructive feeding by the parasitoid larva. 3. Here, a mathematical model is proposed to describe the larval feeding behaviour of the solitary koinobiont endoparasitoid Venturia canescens in four larval stadia of its host Plodia interpunctella. In particular, the model describes how adult size, represented by an exponential growth rate, and development time are traded off when the parasitoid develops in nutritionally suboptimal second stadium hosts. 4. Using a graphical model, the different conditions faced by V. canescens during development in various host species of greatly differing mass are illustrated. 5. It is argued that the relative importance of size and development time on parasitoid fitness is determined by ecological and biological characteristics of both host and parasitoid, and it is suggested that there may be correlations between life-history traits and host-utilisation strategies among koinobionts.     

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。     

Are galling insects better protected against parasitoids than exposed feeders?: a test using tenthredinid sawflies

ABSTRACT.

• 1 Data mostly from the published literature were used to assess the effect of galling on the number of parasitoid species per host species in the phylogeny of nematine sawflies from free external feeders (colonial and solitary) to leaf gallers and shoot gallers.
• 2 The strongest effects of galling were the total elimination of the species-rich cocoon-attacking guild of parasitoids, and eonymphal parasitoids, from the parasitoid community on shoot gallers, all of which are in the genus Euura.
• 3 All tachinid larval parasitoids were also eliminated by the galling habit.
• 4 The cumulative effects of these exclusions resulted in a decline in mean number of parasitoid species per host species from almost sixteen species on external colonial feeders to 4.0 species on shoot gallers.
• 5 General patterns in per cent parasitism by non-tachinid and tachinid larval parasitoids, eonymphal and cocoon parasitoids, on exposed feeders to shoot gallers, showed declines in non-tachinid attack and elimination of tachinid, eonymphal and cocoon parasitoids. But leaf gallers tended to be attacked more than exposed feeders by non-tachinid larval parasitoids.
• 6 The galling habit had a long-term impact by reducing the number of parasitoid species attacking nematine sawfly gallers and per cent mortality inflicted, so that natural enemies may have been important as a selective factor in the evolution of galling nematine sawflies.

     

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.