Grape variety affects female but also male reproductive success in wild European grapevine moths

Abstract.  1. For insect herbivores the quality of the larval host plant is a key determinant of their fitness. Only little attention, however, has been given to the effects of plants on mating success of males and its consequence for the reproductive output of their mates. In addition, almost all the studies that have investigated the influence of host plants on herbivore fitness components have been done in the laboratory, and less is known of these effects in natural conditions.
2. Using the phytophagous European grapevine moth ( Lobesia botrana Den. & Schiff., Lepidoptera: Tortricidae), we tested the influence of grape cultivars as larval food on the probability of acquiring a mate for both sexes, and on the reproductive output of females and males.
3. Results from this study stress the importance of larval host plants on the reproductive success of both sexes. Larval diet differentially affected mating success and reproductive output of male and female moths. Fecundity, egg size, and egg hatchability were significantly different when larvae were fed on particular grape cultivars.
4. A given cultivar that is of poor quality for females is generally also of poor quality for males. A cultivar, however, could be suitable for females but not for males and vice-versa. Apparently, the nutrients required for adult reproduction are not necessarily the same for males and females.
5. The important conclusion from this study is that evaluating the differential effect of host-plant species on traits associated with reproductive success of herbivores requires that the effects on both sexes be taken into account.     

Effects of host variety on blue willow beetle Phratora vulgatissima performance

Phratora vulgatissima adults exhibit feeding preferences among willow varieties, yet little is known of the effects of willow variety on larval and adult performance. The effects of host variety on adult fitness and fecundity as well as on larval mortality and development were studied under laboratory conditions for 35 willow varieties. The host variety significantly affected the number of days that adults survived and the rates of weight change. On average, males lost weight and females gained weight. The total number of eggs, eggs day¹, eggs clutch^?1 and the length of the oviposition period were also significantly affected by willow variety. Progeny from eggs laid by adults fed on the different willow varieties showed significant differences in days to pupation and pupal weight when subsequently reared on Salix × dasyclados or on the same variety as fed to the adults. However, there was no correlation between these parameters on S. × dasyclados and the variety fed to adults. The willow variety fed to larvae significantly affected larval mortality (four varieties caused 100% mortality), the shape of larval growth curves (as measured by predicted final weight and time to half the final weight), the number of days to pupation and pupal weight. There were significant positive correlations between previously determined adult P. vulgatissima feeding preferences of the 35 willow varieties and the following: number of eggs laid, length of the oviposition period, larval mortality and development and change in adult weight. There was a considerable degree of variation in these correlations and some varieties did not follow the general trend indicated by the size or sign of particular correlations, for example, having a high feeding preference ranking yet few eggs laid, low larval weight and longer time to pupation. The differences found between varieties for adult and larval performance in conjunction with previously established feeding preferences offer great potential for utilising plant resistance to P. vulgatissima as a means of strategic control. Despite the general correlation of feeding preference and performance parameters, the results found here suggest that it appears to be possible for plant breeders to circumvent this trend.     

Effects of seed type and bruchid genotype on the performance and oviposition behavior of Zabrotes subfasciatus (Coleoptera: Bruchidae)

The effect of different bean varieties on the performance of the bruchid beetle Zabrotes subfasciatus Boheman (Coleoptera: Bruchidae), was determined by using wild and cultivated seeds of the genus Phaseolus. Results showed that the quality of the host plant affected the performance and the oviposition behavior of female beetles. Overall, bruchid performance was higher on cultivated seeds than on wild seeds. It was also found that the oviposition behavior and the performance of their offspring differed between females that originated from wild versus cultivated seeds. We also demonstrated the importance of a genetic component in bruchid performance: longevity, fecundity, larval development, adult size, and the sex ratio of the progeny varied according to their maternal lineage (‘familyrsquo;). For example, on the same host type, some females laid twice as many eggs as females from other ‘familiesrsquo;, and the sex ratio among the offspring correlated with that of their mother. Thus, the performance and behavior of Z. subfasciatus are not only affected by environmen tal factors such as the quality of the seeds on which they develop, but also have a genetic basis which can counterbalance a less suitable quality of the host plant. For a crop pest such as Z. subfasciatus, its ability to survive and adapt on host plants of differing quality may be an important attribute to consider for pest management.     

Does natural larval parasitism of Lobesia botrana (Lepidoptera: Tortricidae) vary between years, generation, density of the host and vine cultivar?

Populations of European grapevine moth Lobesia botrana Denis & Schiffermüller and its larval parasitoids were studied for two consecutive years on an experimental insecticide-free vineyard in France planted with adjacent plots of five grape cultivars (Merlot, Cabernet Franc, Sauvignon, Cabernet Sauvignon and Sémillon) using a natural L. botrana population during the first year, and a natural population supplemented with artificially inoculated individuals during the second year. Levels of natural populations of larval parasitoids were measured by their parasitism rate. The ichneumonid Campoplex capitator Aubert was the most common species collected from L. botrana larvae. Its incidence was higher during the spring compared to summer. The overall parasitism rate found on the experimental vineyard varied from 23% in 2000 to 53% in 2001, and was mainly due to C. capitator. Parasitism was not affected by the grape cultivar on which the host developed but was positively correlated with the host density, per bunch or per stock, suggesting that among the five grape cultivars tested, C. capitator females probably do not discriminate between hosts feeding on different grape cultivars, but rather the densities of L. botrana larvae.     

Relative performance of European grapevine moth (<Emphasis Type="Italic">Lobesia botrana</Emphasis>) on grapes and other hosts

The European grapevine moth, Lobesia botrana is a major grapevine pest, but despite the abundance of vineyards it is a generalist and uses either grapes or alternative species. Given the abundance and predictability of grape, L. botrana could be expected to have evolved towards monophagy. In order to understand why this species remains polyphagous, we hypothesized that larvae reared on rare wild host plants should have higher fitness than those reared on the more abundant grape host. For this, we compared larval performance and several life history traits on three alternative host plants (Daphne gnidium, Olea europaea, Tanacetum vulgare) and three Vitaceae (Vitis vinifera), two cultivars and one wild species (Ampelopsis brevipedunculata), and two control groups raised on either a low or a high nutritive value medium. Alternative hosts are more suitable than Vitaceae for the reproductive performance of L. botrana: larval mortality and development time was reduced, while pupal weight, growth rate, female longevity, female fecundity, duration of laying and mating success were increased. High quality food ingested by larvae promotes higher adult body weight and enhances female reproductive output. This suggests that alternative hosts provide greater nutritional value for L. botrana than Vitaceae. The use of alternative host plants could thus be maintained in the host range because they offer L. botrana a better fitness than on the Vitaceae. This could typically represent an advantage for moths behaving in plant diversity grape landscapes.     

Greater impact of host plant species on oviposition of a willow leaf beetle <Emphasis Type="Italic">Plagiodera versicolora</Emphasis> Laicharting (Coleoptera: Chrysomelidae) during the adult stage than in developing larval stage

In some herbivorous insects, such as Coleoptera and aphids, not only the host species of larvae, but also those of adults should be considered as key determinants of potential fecundity because oviposition is affected by the quality of host species during both larval and adult stages. This study examined the relatively greater impact on host species of the larval or adult stage on oviposition of the willow leaf beetle Plagiodera versicolora Laicharting (Coleoptera: Chrysomelidae). We conducted an experiment using a 2 × 2 experimental design, in which either of two different host plant species was fed in larval and adult stages. Females fed on a locally unavailable host Salix eriocarpa in the adult stage did not lay any eggs, but those fed on the locally available host S. babylonica laid 67–75 eggs on average, irrespective of larval host species. Such reproductively inactive females fed S. eriocarpa as an adult host recovered reproductive activity within 3 weeks after changing the host species to S. babylonica. This result indicated that the host species fed in the adult stage had a greater impact on oviposition than in the larval stage.     

Feeding insects with a phytopathogenic fungus influences their diapause and population dynamics

Abstract.  1. A facultative mutualistic relationship between a herbivorous moth, Lobesia botrana , and a phytopathogenic fungus, Botrytis cinerea , both damaging the same plant, the vine, has been demonstrated recently. Laboratory and field studies were carried out to determine the influence of the presence of the fungus in food on larvae during the pre- and post-diapause periods and on the global fitness of the insect.
2. During the pre-diapause period, larvae fed on fungus exhibited a higher survival rate and a faster larval development than insects reared without fungus.
3. After the wintering diapause, insects reared with fungus displayed a better synchronisation of adult emergence and an increased fecundity compared with control insects.
4. These results indicate that food containing fungus provides an improved diet, permitting the herbivorous insect to obtain nutrients required to optimise their life-history traits.
5. In addition, the results emphasise the importance of the nutritional quality of larval diet, improved by addition of the fungus, on diapause termination and on the demography of the insect before and after the wintering diapause.     

Host plant exploitation within a population of a generalist herbivore, Choristoneura rosaceana

Feeding performance of larvae of the obliquebanded leafroller, Choristoneura rosaceana (Harris), was estimated on apple and three alternative hosts in the Okanagan Valley. The obliquebanded leafroller is native to the study area and apple was introduced in the valley at the beginning of the century. Components of insect fitness were measured to help formulate predictions about the evolution of host preference in this generalist herbivore. In the summer, females attained a higher pupal weight on apple than on trembling aspen, wild rose and snowberry. Males performed best on apple and trembling aspen. In the field, females arising from caterpillars fed on apple leaves had a 26% greater mean fecundity than if fed on alternative hosts. The diet also affected development time; females initiated pupation later on trembling aspen than on the other hosts. Finally, no differences were found in larval survival nor in the proportion of larvae diapausing when fed on different hosts. These components of fitness indicate apple as the most suitable host in the summer.In the fall, diet influenced larval propensity to initiate diapause in the second or third instar, the time elapsed from hatching to diapause initiation and the weight of the instars initiating diapause. However, no difference in overwintering survival among larvae fed on different hosts was detected. The rank order of water and nitrogen contents of leaves of the host plants did not match the rank order of larval feeding performance in either season. A high suitability of apple and its abundance in the Okanagan valley are factors that would favor the evolution of an increased utilization of this host by the obliquebanded leafroller.     

外来植食性广聚萤叶甲对非靶标植物的潜在影响

为评估以入侵豚草为食的外来广聚萤叶甲（Ophraella communa）对非靶标植物的潜在生态风险,采用选择性试验测定了广聚萤叶甲成虫和幼虫对当地代表性植物的取食选择,观察了幼虫在选出的植物上生长发育的适合度表现。结果表明：在5大类52种测试植物中,广聚萤叶甲成虫和幼虫不同程度地取食向日葵、苍耳、天明精、菊芋（仅成虫）、紫茎泽兰（仅幼虫）和石胡荽（仅幼虫）;野外罩笼非选择性测定发现,初孵幼虫在苍耳和向日葵8个品种上可完成发育,直到成虫产卵,并观察到在菊芋上完成幼虫发育,个别到成虫但未产卵;在这些植物上幼虫存活率明显降低,蛹质量明显减轻,成虫产卵明显减少。本文还分析了广聚萤叶甲对少数本土植物以及经济作物向日葵的潜在生态风险。     

Stage-specific control of grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortricidae), by selective and broad-spectrum insecticides

The insect growth regulators (IGRs) tebufenozide and methoxyfenozide and the broad-spectrum insecticides azinphosmethyl, carbaryl, and fenpropathrin were compared for their activity against adult, egg, and larval stages of the grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortricidae), under laboratory and vineyard conditions. Adult mortality was not affected by exposure to field-equivalent rates of tebufenozide or methoxyfenozide on grape clusters, whereas all the broad-spectrum compounds significantly reduced adult survival, compared with the untreated controls. Surviving adult moths laid significantly more eggs on berries treated with the IGRs than on berries treated with any of the broad-spectrum insecticides. Survival of these eggs through to late larval and pupal stages was significantly lower on methoxyfenozide-treated grapes than on untreated grapes, and no pupae were found when grapes were treated with azinphosmethyl or fenpropathrin. Neither of the growth regulator insecticides limited egg eclosion or larval development by E. viteana when insecticides were applied before egg laying, whereas broad-spectrum insecticides were effective against both eggs and neonates at this timing. When applied after egg eclosion, all insecticide treatments significantly reduced survival of grape berry moth larvae. Under vineyard conditions, berries with 1-d-old residues of tebufenozide or methoxyfenozide received more E. viteana eggs than berries treated with broad-spectrum compounds. After aging for 7 or 14 d, no significant effects on E. viteana survival were detected among treatments. Whereas broad-spectrum insecticides provide control of multiple life stages of E. viteana, integration of tebufenozide or methoxyfenozide into vineyard management programs for control of this pest will be most successful if applications are timed for egg hatch.     

Egg size variation and its relationship with larval performance in the Lepidoptera: the case of the European grapevine moth Lobesia botrana

Life history theory attempts to define the pattern of resource distribution among offspring and predicts that egg size should be positively correlated with offspring fitness. In this paper we investigated the effect of an array of ecological and reproductive factors on the size of eggs laid by Lobesia botrana, and the ecological significance of egg size by experimentally testing whether or not egg size increased larval performance in this moth. Egg size was significantly affected by female age at mating, water availability, pupal weight loss, the phenological stage of the vine in which larvae developed, female body weight, and oviposition day, but was unaffected by the size of the spermatophore received. The greater the size of the first eggs produced, the smaller the size of the last eggs laid, as predicted by the resource depletion hypothesis. As expected, the larger the size of eggs the larger the size of larvae emerging from them. Large larvae displayed a better ability to endure starvation than small ones. When small and large larvae were allowed to develop on grape clusters, an adverse environment, large larvae performed better, settling and survival being significantly enhanced, almost tripled. Instead, when the same experiment was carried out on an artificial diet in the laboratory, a much more favourable milieu, survival in both sized-larval groups was not significantly different. We discuss our results in the context of current ideas relating egg size, larval performance and fitness in the Lepidoptera.     

Differences in nutritional quality of parts of Vitis vinifera berries affect fitness of the European grapevine moth

The European grapevine moth, Lobesia botrana (Denis & Schiffermüller) (Lepidoptera: Tortricidae), is a major grapevine pest in Europe. The larva is polyphagous and able to develop on more than 25 plant species, several of them being more suitable than Vitaceae for the fitness of L. botrana. Larvae normally eat the pulp of the berry, but may also consume the seeds according to the development stage of the berry and the larval density per bunch. Understanding the effect on individual fitness of such feeding behaviour is important to assess how suitable the different berry tissues are for this insect. We offered to the larvae either entire berries, seeds, or pulp with skin of the variety Vitis vinifera cv. ‘Cabernet Sauvignon’ as larval food in order to assess several life history traits from egg hatching to adult death. Two control groups were raised on semisynthetic diets, offering a low (no plant material) or a higher (with corn flour) nutritive value medium. The larvae performed differently when fed on diets containing different parts of berries. The larvae fed on a diet containing grape berry seeds had a prolonged development time and showed higher mortality. The females emerging from these larvae had a lower fecundity and mating success than the females emerging from larvae fed on diets containing other parts of the berries. However, their longevity was greater in comparison to the other groups. We conclude that seeds of Cabernet Sauvignon are not toxic to larvae but affect the reproductive life history traits in L. botrana.     

Host selection in phytophagous insects: a new explanation for learning in adults

Insect learning can change the preferences an egg laying female displays towards different host plant species. Current hypotheses propose that learning may be advantageous in adult host selection behaviour through improved recognition, accuracy or selectivity in foraging. In this paper, we present a hypothesis for when learning can be advantageous without such improvements in adult host foraging. Specifically, that learning can be an advantageous strategy for egg laying females when larvae must feed on more than one plant in order to complete development, if the fitness of larvae is reduced when they switch to a different host species. Here, larvae benefit from developing on the most abundant host species, which is the most likely choice of host for an adult insect which increases its preference for a host species through learning. The hypothesis is formalised with a mathematical model and we provide evidence from studies on the behavioural ecology of a number of insect species which demonstrate that the assumptions of this hypothesis may frequently be fulfilled in nature. We discuss how multiple mechanisms may convey advantages in insect learning and that benefits to larval development, which have so far been overlooked, should be considered in explanations for the widespread occurrence of learning.     

Butterfly oviposition preference is not related to larval performance on a polyploid herb

The preference–performance hypothesis predicts that female insects maximize their fitness by utilizing host plants which are associated with high larval performance. Still, studies with several insect species have failed to find a positive correlation between oviposition preference and larval performance. In the present study, we experimentally investigated the relationship between oviposition preferences and larval performance in the butterfly Anthocharis cardamines. Preferences were assessed using both cage experiments and field data on the proportion of host plant individuals utilized in natural populations. Larval performance was experimentally investigated using larvae descending from 419 oviposition events by 21 females on plants from 51 populations of two ploidy types of the perennial herb Cardamine pratensis. Neither ploidy type nor population identity influenced egg survival or larval development, but increased plant inflorescence size resulted in a larger final larval size. There was no correlation between female oviposition preference and egg survival or larval development under controlled conditions. Moreover, variation in larval performance among populations under controlled conditions was not correlated with the proportion of host plants utilized in the field. Lastly, first instar larvae added to plants rejected for oviposition by butterfly females during the preference experiment performed equally well as larvae growing on plants chosen for oviposition. The lack of a correlation between larval performance and oviposition preference for A. cardamines under both experimental and natural settings suggests that female host choice does not maximize the fitness of the individual offspring.     

Polyphagy,flightlessness, and reproductive output of females: a case study with bagworms (Lepidoptera: Psychidae)

Abstract 1. We evaluated the relation between plant‐mediated larval traits and the fitness of female bagworms, Thyridopteryx ephemeraeformis Haworth (Lepidoptera: Psychidae) by sampling 29 populations of bagworms on five plant genera (Thuja, Pinus, Picea, Juniperus, and Gleditsia). 2. Compared with those on other genera of plants, female larvae on Thuja attained a larger size at pupation and developed faster. The superior quality of Thuja as a larval food plant resulted in a higher potential fitness of females, as indicated by a low level of pupal mortality and high fecundity. 3. The fecundity of females increased with pupal size on different plant genera, and the effect of host plant genus was negligible compared with pupal size. 4. The mating success of females varied among different populations (0–37% unmated females), but neither host plant, emergence time, nor the size of females had a significant influence on the proportion of mated females. 5. The conversion of adult biomass into reproductive tissue, measured as the ratio between the biomass of eggs divided by the biomass of calling females, increased with the pupal size of females and approached 90% for large females. The high rate of egg conversion in bagworms may be related to the neoteny of short‐lived females that invest little in somatic tissue.     

Oviposition preference,larval performance and adaptation of Trichoplusia ni on cabbage and cotton

Most female herbivores ensure to lay eggs where their offspring can develop successfully. The oviposition preferences of females affect strategies in pest management. In this study, the performance of two cohorts of Trichoplusia ni larvae on cabbage and cotton (after they had been transferred from their original host plants) were investigated. The preferences of female moth ovipositing and larval feeding on these two host plants were observed. The results indicated that plants significantly affected oviposition preference of the female adults and development and survival of larvae of T. ni. All females preferred to lay eggs on cabbage than cotton regardless from which host they originated. The detrimental effects of cotton on the development and survival of T. ni larvae originated from cabbage (CaTn) increased with the increase of the larval age when they were transferred. In addition, the host plant change did not significantly affect the development and survival of larvae of T. ni originating from cotton (CoTn). Larvae of CaTn preferred cabbage plants as compared to cotton plants, whereas larvae of CoTn did not show a significant choice. Although the adult females preferred laying eggs on cabbage, they did not show preferences between cotton and cabbage in a Y‐tube olfactometer test. The hypothesis of oviposition preference and performance of larvae was supported by the results of CaTn, whereas they not supported by those from CoTn. Based on these results, the strategy to manage this serious pest was discussed.     

Oviposition,larval preference,and larval performance in two polyphagous species: does the larva know best?

It is expected that females preferentially oviposit on plant hosts that allow for optimal larval performance. However, this expectation contradicts empirical evidence where adults do not always choose the best host for their descendants. Recent evidence suggests that females’ host selection depends on the number of potential hosts. Females from oligophagous species seem to be able to choose an appropriate host in terms of larval performance, whereas in polyphagous species, adult oviposition preference is not related with larval performance. This suggests that larvae in polyphagous species could be taking a more active role in host selection than their mothers. Here, we evaluated the oviposition preference and the larval preference and performance of two polyphagous species of economic importance, Copitarsia decolora (Guenée) (Lepidoptera: Noctuidae: Cuculliinae) and Peridroma saucia (Hübner) (Lepidoptera: Noctuidae: Noctuinae), on eight species of cultivated plants. In laboratory and greenhouse choice assays, we tested adult preference for oviposition and larval preference at 1 and 24 h. Larval performance was measured in terms of survival to adulthood, length of larval period, and pupal weight. We found that both adult females and larvae actively choose their hosts and that the larval preference toward the hosts is related to the females’ preference in both herbivore species. However, the females and larvae did not preferentially select the host with the best larval performance, indicating that larval performance is not related to female or larval preference and that other selective pressures are influencing the choice of the host plant in these two species.     

Relationship between adult and larval host plant selection and larval performance in the generalist moth, Trichoplusia ni

Adult oviposition preferences are expected to correlate with host plant suitability for the development of their offspring. For most lepidopteran species, this is particularly important as the hatching neonate larvae of many species are relatively immobile. Thus, the site of oviposition chosen by a female adult can greatly influence the probability of survival for her offspring. In the present study, we investigated the oviposition preference of adult Trichoplusia ni moths for six plant species to determine whether they could accurately rank the suitability of the plants for larval development. We also compared oviposition preferences to neonate larval acceptance and preference to determine whether the adult host range matched that of larval diet breath. Our results indicate that in two-choice and no-choice tests adult T. ni were able to rank the plants accurately, with the exception of anise hyssop. However, when given a choice of all six plants together, they laid more eggs on a plant that was not suitable for larval survival. Larvae accepted and fed on all plants in no-choice tests, and accurately ranked them according to larval performance. We conclude that neonate larvae are better able than adults to rank plants according to larval performance, and that larval diet breadth is wider than the range of plants accepted by adults. We also provide a discussion of the reduced accuracy of adult oviposition preference with increased plant choices.     

Susceptibility of cold hardy grapes to Drosophila suzukii (Diptera: Drosophilidae)

Drosophila suzukii (Diptera: Drosophilidae), known commonly as spotted wing drosophila, is a vinegar fly originating from South‐East Asia and a major pest to many soft‐skinned fruits. Due to the species recent arrival in North America in 2008, many fruit varieties are yet untested for susceptibility to infestation. While previous work has focused on Vitis vinifera, this study aimed to determine grape susceptibility of cold hardy varieties based on hybrids of V. labrusca, V. riparia and V. vinifera. Field sampling was conducted in Southern Wisconsin (USA) vineyards to establish adult and larval abundance and determine whether the number of adults caught in traps correlates with fruit infestation. Host susceptibility was further assessed through no‐choice bioassays of both intact and damaged fruits. The field study found D. suzukii adults present in all varieties, low larval abundance and no correlation between adult abundance and larval presence. Peak adult abundance occurred mid‐season between veraison and harvest, while larval infestation rates were highest near harvest. In laboratory no‐choice tests, significantly more eggs, larvae and adults occurred in damaged than undamaged grapes. In damaged grapes, larvae and adult abundance was comparable between varieties and to the highly susceptible control of undamaged raspberry; however, D. suzukii developed significantly faster in raspberry than grapes. Fruit characteristics (°Brix, titratable acidity, pH) in grapes were uncorrelated with D. suzukii performance. Together, these findings suggest that cold hardy grapes are overall resistant to D. suzukii if intact and highly susceptible if damaged.