Temperature-induced changes in spider mite fitness: offsetting effects of development time, fecundity, and sex ratio

Temperature affects many life history parameters in poikilotherms. Temperature clearly affects development time and fecundity, which affect the intrinsic rate of increase. In haplodiploid mites, ambient temperature may also affect offspring sex ratio which, in turn, affects intrinsic rate of increase. The combined effect of all these processes determines the fitness of individual females. However, sex ratio also affects mating structure and, potentially, rate of local adaption. We investigated the direct effect of temperature variation on sex ratio, development time, and fecundity in the twospotted spider mite (Tetyranychus urticae), and calculated the effect of their interaction on mite intrinsic rate of increase. We conducted experiments at 2 temperatures and designed the experiment to separate pre-adult and adult sensitivity to temperature variation. Mites were reared from eggs to adult ecdysis at either 22°C or 32°C. Upon emergence as adults, these 2 groups were each split between 22°C and 32°C and allowed to oviposit. Not surprisingly, development from egg to adult was accelerated when mites were exposed to the higher temperature during offspring development, regardless of the temperature experienced by the mother during her development. Fecundity and the proportion of female offspring were affected by temperature only when mothers were exposed during both development and oviposition. About 12 offspring were gained and female bias was increased by 26% when the mother's development occurred at 22°C, whereas oviposition at 22°C added only 6 more offspring and increased female bias by only 7%. There was no correlation between sex ratio and fecundity; both were related to temperature but not to each other. Furthermore, development time, not fecundity or sex ratio, appeared to the main factor affecting the intrinsic rate of increase. Our results support other evidence that sex ratio varies independently of development time and fecundity.     

寄主体型大小对管氏肿腿蜂生殖潜力的影响

【目的】揭示寄主体型大小对肿腿蜂生殖潜力的影响,进一步认识寄生蜂准社会行为的进化以及为大规模繁殖和利用肿腿蜂的生物防治实践提供科学依据。【方法】以管氏肿腿蜂Sclerodermus guani和寄主青杨天牛Saperda populnea幼虫为材料,为雌蜂终身连续提供称重的寄主,观察逐日产卵量及在无抚育条件下其子代蜂发育至成虫的表现,分析产卵时期(早、中和晚期)、寄主体重对产卵量及其子代发育指标的影响。【结果】寄主体重和产卵期均影响管氏肿腿蜂终身产卵量,产卵量随寄主体重增大而呈二次函数式增大,但随产卵期延长而大幅减小。寄主体重和产卵期互作影响管氏肿腿蜂子代蜂存活率、岀蜂数、成虫体重和性比等子代发育指标,表现在:(1)产卵早期所产子代蜂的存活率随寄主体重增大而略降低,产卵中期所产子代蜂存活与寄主体重无关,产卵晚期所产子代蜂的存活率可能与寄主体重无关;(2)产卵早期和晚期产子代蜂的羽化成虫数和性比不受寄主体重的影响,但产卵中期产的子代蜂出蜂数随寄主体重增大而增加,性比随寄主体重增大而逐渐减小;(3)产卵早期产的子代雌蜂体重随寄主体重增大而呈二次函数式增大,而产卵中期产的子代蜂体重不受寄主体重的影响。【结论】寄主体型大小对管氏肿腿蜂早期产卵量及其子代蜂发育表现有显著影响。     

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.     

Functional response and mutual interference of Peristenus spretus (Hymenoptera: Braconidae), a parasitoid of Apolygus lucorum (Heteroptera: Miridae)

Peristenus spretus Chen & van Achterberg (Hymenoptera: Braconidae) is a solitary endoparasitoid, which is considered for augmentative biological control of Apolygus lucorum Meyer-Dür (Heteroptera: Miridae) in Chinese cotton fields. Since the association of P. spretus with A. lucorum was only recently discovered, the biology of the parasitoid remains unknown. In order to understand its reproductive biology, the mutual interference and functional response of P. spretus were investigated by altering either the parasitoid or the host density while keeping the other constant. In both experiments, the effects of parasitoid and host densities on parasitism, superparasitism, progeny production and sex ratio were assessed. P. spretus exhibited a Holling type II functional response to changing host densities, indicating that parasitism increases with increasing host density until the parasitoid reaches its maximum reproductive capacity. The model suggested that a single P. spretus female could parasitise a maximum of 88 nymphs per day or four nymphs per hour. Increasing the wasp-nymph ratio from 1:10 to 1:80 significantly increased the offspring production more than fivefold from ±5.8 to ±35.6; further increasing the host densities (above 80 nymphs) did not significantly increase offspring production. Strong mutual interference of foraging P. spretus females occurred only at high parasitoid densities. Parasitoids foraging alone produced an average progeny of 33.4, whereas parasitoids foraging in groups of 16 produced only 2.6. The optimal wasp-nymph ratio for mass-rearing P. spretus is 4:100, given that resources of parasitoids and nymphs are unlimited.     

Sex allocation and egg distribution of an autoparasitoid, Encarsia pergandiella (Hymenoptera: Aphelinidae)

Abstract. 1. Encarsia pergandiella Howard females develop as primary parasitoids on immature whiteflies, and males develop as secondary parasitoids on females of their own or a related species. The hypothesis that the sex ratio reflects the relative abundance of the two host types was tested in the laboratory using petri dish arenas with varying proportions of early fourth instar greenhouse whitefly (Trialeurodes vaporariorum (West.)) (primary hosts) and pupal female E.pergandiella (secondary hosts). Egg distribution was analysed with respect to sex ratio, super-parasitism and host discrimination.
2. The proportion of primary and secondary hosts parasitized in each treatment reflected the relative availability of each host type. Thus females presented with 75% primary hosts laid more female eggs than male. However, in all treatments, a greater proportion of secondary hosts were parasitized than would be expected from the proportion of secondary hosts available. This indicates that more male eggs were laid than expected.
3. More secondary hosts than primary hosts were superparasitized.
4. Host discrimination analysis using a new test statistic showed that females generally laid eggs at random with regard to previous parasitism of primary or secondary hosts. However, females in one treatment with 50% of each host type appeared to preferentially oviposit in secondary hosts which did not contain any eggs.     

Reproductive Biology ofCephalonomia hyalinipennis(Hymenoptera: Bethylidae), a Native Parasitoid of the Coffee Berry Borer,Hypothenemus hampei(Coleoptera: Scolytidae), in Chiapas, Mexico

The coffee berry borer,Hypothenemus hampei(Ferrari) (Coleoptera: Scolytidae), is the most important pest of coffee worldwide.Cephalonomia hyalinipennisAshmead (Hymenoptera: Bethylidae) was recently discovered naturally attackingH. hampeiin Mexico.C. hyalinipennisboth preys upon and parasitizesH. hampei.We report laboratory investigations on the reproductive biology ofC. hyalinipennisaimed at assessing the suitability of this parasitoid for culturing and release in biological control programs. Adult females lived up to 95 days (mean for mated females = 57 days), and risk of death increased with age. Mating status, reproductive effort, and female size influenced adult female longevity, but only weakly. The number ofH. hampeipreyed upon did not affect longevity. Rates of attack (predation plus parasitism) declined with age, but the proportion of attackedH. hampeithat were parasitized increased. Overall, about half of the attackedH. hampeiwere preyed upon and half were parasitized. Females received approximately 46 sperm per mating and 77% became sperm depleted before the end of their reproductive lives. Estimated mean lifetime fecundity was 88 eggs. Offspring survival from egg to adulthood was 60%, with batches of eggs tending to survive or die collectively; 21.2% of surviving progeny were males. These results suggest thatC. hyalinipennismay be suitable for mass rearing and release, but further work is needed to investigate interactions betweenC. hyalinipennisand other parasitoids ofH. hampei.     

The influence of a shortage of hosts on the specificity of oviposition behaviour in species of Dacus (Diptera, Tephritidae)

ABSTRACT. When deprived of their preferred hosts for up to 16 days prior to behaviour assays, three relatively specialized tephritid flies, D. cacuminatus (Hering), D. cucumis French and D. jarvisi (Tryon), retain strong preferences for their particular hosts. Moreover, in the absence of the preferred hosts, D. cacuminatus , and to a lesser extent D. cucumis , will not accept fruits that are outside their normal host range. D. jarvisi readily accepts other fruits in the absence of its preferred host, Planchonia careya , but in common with the other two shows no increase in the number of eggs laid with increasing periods of deprivation. None of these species accumulate oocytes when deprived of fruits.
In contrast, the highly polyphagous D. tryoni Frogg. lays more eggs and oviposits readily into a previously unacceptable fruit after 4 days without access to hosts. This change in behaviour is associated with a marked increase in the number of mature eggs carried by females.
These results suggest an important difference between the generalist and more specialized species in the physiological control of oocyte maturation. In D. tryoni there appears to be no inhibition of oocyte development once the primary follicle has matured, whereas in the other species each ovariole contains no more than one mature egg.
In D. tryoni , the change in behaviour which accompanies this increase in egg load, may influence its propensity to accept unusual fruits, and may help to explain its rapid acceptance of most cultivated fruits introduced to Australia.     

Life history and host range of Alagoasa extrema (Coleoptera: Chrysomelidae: Alticinae), a potential biological control agent of Lantana spp. (Verbenaceae) in Australia

The life history and host range of the lantana beetle, Alagoasa extrema, a potential biocontrol agent for Lantana spp. were investigated in a quarantine unit at the Alan Fletcher Research Station, Brisbane, Australia. Adults feed on leaves and females lay batches of about 17 eggs on the soil surface around the stems of plants. The eggs take 16 days to hatch and newly emerged larvae move up the stem to feed on young leaves. Larvae feed for about 23 days and there are three instars. There is a prepupal non-feeding stage that lasts about 12 days and the pupal stage, which occurs in a cocoon in the soil, lasts 16 days. Teneral adults remain in the cocoon for 3 days to harden prior to emergence. Males live for about 151 days while females live for about 127 days. The pre-oviposition period is 19 days. In no-choice larval feeding trials, nine plant species, representing three families, supported development to adult. Three species, Aloysia triphylla, Citharexylum spinosum and Pandorea pandorana were able to support at least two successive generations. These results confirm those reported in South Africa and suggest that A. extrema is not sufficiently specific for release in Australia. Furthermore, it is not recommended for release in any other country which is considering biological control of lantana.     

COI and ITS2 sequences delimit species, reveal cryptic taxa and host specificity of fig-associated Sycophila (Hymenoptera, Eurytomidae)

Although the genus Sycophila has broad host preferences, some species are specifically associated with figs as nonpollinator wasps. Because of their sexual dimorphism, morphological plasticity, cryptic mating behaviour and poorly known biology, species identifications are often uncertain. It is particularly difficult to match conspecific females and males. In this study, we employed two molecular markers, mitochondrial COI and nuclear ITS2, to identify Sycophila from six Chinese fig species. Morphological studies revealed 25 female and male morphs, while sequence results for both genes were consistent in supporting the presence of 15 species, of which 13 were host specialists and two used dual hosts. A single species of Sycophila was respectively found on four fig species, but six species were isolated from Ficus benjamina and a same number was reared from Ficus microcarpa. Sequence results revealed three male morphs in one species and detected two species that were overlooked by morphological analysis.     

辣椒碱对烟粉虱的生物活性及生长发育和生殖力的影响

在实验室条件下研究了辣椒碱对烟粉虱的生物活性及生长发育和生殖力的影响.结果表明:辣椒碱对烟粉虱各虫态均有杀虫活性,且随辣椒碱浓度的增大活性增强;辣椒碱浸叶法/浸虫法处理对烟粉虱成虫、卵和若虫的毒力(LC50分别为3525.79、1603.29和1526.27mg·L-1))均高于内吸法处理(LC50分别为5360.04、2359.19和5897.43 mg.L-1),两种处理方法下辣椒碱对烟粉虱卵的毒力均明显高于对成虫的毒力;2000和4000 mg·L-1浓度辣椒碱对烟粉虱成虫具有明显的拒食作用,烟粉虱在该两种浓度辣椒碱处理过的棉花叶上取食24 h的蜜露分泌量均显著低于取食非处理棉花叶后的蜜露分泌量;辣椒碱各试验浓度对烟粉虱成虫均表现出较强的产卵忌避作用,当辣椒碱处理浓度达4000 mg·L-1时,对烟粉虱的选择性产卵忌避率和非选择性产卵忌避率分别高达94.2％和83.0％.500、1000和2000 mg·L-1辣椒碱浸渍处理对烟粉虱存活卵和若虫的发育历期、成虫羽化率均没有影响,但辣椒碱4000mg·L-1处理明显延长烟粉虱若虫发育历期、降低成虫羽化率.不同浓度辣椒碱处理均明显抑制烟粉虱生殖力,且这种抑制作用随辣椒碱浓度的增大而增强.     

The effect of superparasitism on development of the solitary parasitoid wasp, Venturia canescens (Hymenoptera: Ichneumonidae)

Abstract.

• 1 The effects of superparasitism on the rate of development, adult size and mortality of Venturia canescens (Hymenoptera: Ichneumonidae) were investigated. Parasitoids were reared from third (L3) and fifth (L5) instars of one of its hosts, Plodia interpunctella (Lepidoptera: Pyralidae) containing one, two or four parasitoid eggs.
• 2 Superparasitism increased the development time of Venturia reared from both instars, but the developmental delay was more evident in wasps from L5 hosts.
• 3 The size of parasitoids from L3 hosts was unaffected by egg number, but wasps from both superparasitized L5 treatments were significantly smaller than those from singly parasitized hosts.
• 4 Parasitoid mortality was significantly higher in L5 than L3, but within instars did not differ significantly with egg number.
• 5 The results confirm that superparasitism may affect the fitness of both the adult female wasp and her progeny, and should therefore be incorporated into models of superparasitism as an adaptive foraging strategy.

     

Effects of temperature on development,survival, longevity,and fecundity of Serangium japonicum (Coleoptera: Coccinellidae), a predator of Bemisia tabaci Gennadius (Homoptera: Aleyrodidae)

In this study, we evaluated the effect of temperature on the development and reproductive biology of Serangium japonicum (Coleoptera: Coccinellidae) at seven constant temperature regimes (17, 20, 23, 26, 29, 32 and 35°C) for its effect as a predator of Bemisia tabaci (Homoptera: Aleyrodidae). Results indicated that the duration of the egg, larval and pupal stages were significantly affected by temperature. The developmental time gradually declined with the increase of temperature from 17 to 29°C, however an extension in the developmental periods was observed in the temperature range of 32 to 35°C. The survival rates of different insect stages were stable at temperatures between 20 and 32°C; however at extreme temperatures of 35°C, a sharp decrease was evident. The highest fecundity of the female (387.2 eggs per female) was recorded at 20°C. Based on these results, life tables of S. japonicum were constructed for temperatures in the range 20–35°C. The maximum reproductive rate (R ₀=279.9) occurred at 26°C. The maximum values for innate capacity for increase (r _m=0.1131) and the finite rate of increase (λ=1.1197) occurred at 29°C. The mean generation time (T) decreased with increased temperature, the longest of which was 76.0 days (at 20°C) and the shortest was 36.6 days (at 32°C). These results offer valuable insight on the importation and establishment of S. japonicum into new environments with diverse temperature regimes.     

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.     

Suitability of various stages of mealybug,Phenacoccus solenopsis (Homoptera: Pseudococcidae) for development and survival of the solitary endoparasitoid,Aenasius bambawalei (Hymenoptera: Encyrtidae)

Aenasius bambawalei has been recently reported as a solitary endoparasitoid of Phenacoccus solenopsis in India. Laboratory experiments were conducted comparing the parasitoid's preference for the three developmental stages of mealybug and its effect on parasitism, development, progeny fitness and sex ratio of A. bambawalei. The studies revealed third instar mealybugs as the most preferred stage for development of the parasitoid.     

Variable sex ratio strategy of Telenomus heliothidis (Hymenoptera: Scelionidae): adaptation to host and conspecific density

Summary The sex allocation behavior of the solitary egg parasitoid Telenomus heliothidis Ashmead was investigated by examining the response of females reared in isolation and under crowded conditions. Females reared in isolation adjusted their sex ratio with foundress and host number per patch in accordance with the predictions of local mate competition (LMC) theory. However, females did not shift their sex ratio in response to conspecifics foraging on the same host patch or to contact with previously parasitized hosts. Instead, shifts were associated with encounter rate and a sequence of oviposition. Females maintained under crowded conditions responded to host patches much differently. One-day-old females which had lived under crowded conditions for 24 h produced sex ratios similar to those of continuously isolated females. However, females reared under crowded conditions for 7 days consistently produced unbiased sex ratios, and exhibited a different sequence of oviposition. This shift appeared to be due directly to crowding rather than age, oviposition experience or sperm depletion since the effect could be reversed by subsequent isolation.     

Effect of temperature on the reproductive biology of Peristenus spretus (Hymenoptera: Braconidae), a biological control agent of the plant bug Apolygus lucorum (Hemiptera: Miridae)

Peristenus spretus Chen et van Achterberg (Hymenoptera: Braconidae), a parasitoid of the plant bug Apolygus lucorum (Hemiptera: Miridae), has been studied for use in augmentative biological control in China. Under laboratory conditions, we explored the development, survival, age-specific and potential lifetime fecundity, oviposition period and progeny sex ratio of P. spretus reared at six constant temperatures (15°C, 19°C, 23°C, 27°C, 31°C, 35°C) on the second instar nymphs of A. lucorum. At 15°C, male and female P. spretus took 48.7 ± 0.3 and 52.5 ± 0.3 days to complete their immature development, while developmental time was reduced by more than half at 23°C and 27°C. The parasitoid can only develop to the larval stage at 31°C and neither larva nor pupa survived at 35°C. The estimated lower developmental threshold of the immature stage was 7.3°C. When parasitoid adults were exposed at 15°C, females laid 90% of their eggs at first 19 days of oviposition and had an extended reproductive life. In contrast, females held at 27°C laid most of their eggs (90%) in their first of 10 days of oviposition and had shorter longevity. The highest potential lifetime fecundity of P. spretus was 671.2 ± 34.7 SE eggs produced over 23.4 ± 1.4 SE days at 23°C. At 15°C, 19°C and 23°C, sex ratios of reared parasitoids were male-biased, but at 27°C there was no male bias.     

Self-, Intra-, and Interspecific Host Discrimination in Telenomus busseolae Gahan and T. isis Polaszek (Hymenoptera: Scelionidae), Sympatric Egg Parasitoids of the African Cereal Stem Borer Sesamia calamistis Hampson (Lepidoptera: Noctuidae)

Oviposition behavior, intra- and interspecific host discrimination, and super- and multiparasitism by the scelionids Telenomus busseolae and T. isis were studied using batches of eggs of the noctuid stalk borer Sesamia calamistis as the host. Both Telenomus species were able to discern eggs already parasitized. As a result, self-superparasitism was only 4.0% for T. busseolae and 5.8% for T. isis. Likewise, intraspecific superparasitism was avoided by both species and was significantly higher for T. busseolae than T. isis, and higher when parasitized eggs were offered immediately (0 h) and after 48 h than after 24 h; apparently, the recognition of parasitized eggs at 24 and 48 h was based on the presence of parasitoid larvae rather than a specific marking substance. Multiparasitism was avoided if the female had a choice between unparasitized and parasitized eggs. In a choice experiment, it was 10.2 and 2.5% for T. busseolae and T. isis, respectively. In a nonchoice experiment, multiparasitism did not vary between species and time treatment; it was low, varying between 9.6 and 24.1%. In the 0-h treatment, T. busseolae always outcompeted T. isis, accounting for 63.4 or 91.7% of the offspring, when T. isis or T. busseolae, respectively, was the first species. In the 24-h treatment, the first species emerged more often than the second did but the host egg mortality, i.e., eggs from which neither borer larvae nor parasitoid emerged, was >40%. Whereas T. busseolae is ubiquitous in Africa, T. isis has never been reported from eastern Africa, and it has been proposed for introduction against the prevailing noctuid pest Bussoela fusca. Based on earlier life table and host suitability studies and on the present findings, it is concluded that T. isis would establish in the midaltitudes but not the highlands of eastern Africa.     

The influence of host size on sex ratios in the parasitoid Diglyphus begini (Hymenoptera: Eulophidae)

Abstract. 1. Sex ratio in the ectoparasitoid, Diglyphus begini (Ashmead), attacking larvae of the dipterous leafminer Liriomyza trifolii (Burgess) in glasshouse marigolds, is best depicted by a model where sex ratio varies as a function of host size.
2. Male D. begini progeny are produced in hosts significantly smaller in size than those producing female progeny.
3. Female wasps attack and oviposit on the largest leafminer larvae available and whether a host is large or small depends upon the size of the other hosts attacked.
4. Diglyphus begini females adjust the thre:shold size for the change-over in sex allocation relative to the size of hosts attacked; however, the sex ratio is maintained at between 60% and 70% males.
5. The patterns observed in these glasshouse studies are not due to sex-specific differential mortality or superparasitism.     

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16′ S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.     

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.