Biotypes of Dasineura tetensi, differing in ability to gall and develop on black currant genotypes

Variation in damage levels on certain black currant, Ribes nigrum L., genotypes, caused by the black currant leaf midge, Dasineura tetensi (Rübs.) (Diptera: Cecidomyiidae), has been observed in northern Sweden. I investigated whether this variation is due to variation in virulence among midges. From a field population of midges, I successfully selected for virulence and avirulence, respectively, on the resistant black currant genotype cultivar `Storklas' (called resistant genotype). The performance of avirulent and virulent midge larvae on two black currant genotypes were studied in experiments where first or second instar larvae were artificially transferred. There were no differences in larval survival and developmental rate between the two midge types when transferred to the susceptible currant genotype `7801–31' (called susceptible genotype). Larvae of the virulent strain established galls and developed on `Storklas' but development was initially slower there than on the susceptible currant genotype. Larvae of the avirulent strain suffered high mortality or remained in first instar on that same currant genotype when transferred alone, but developed readily if transferred together with virulent larvae. Larvae transferred in second instar to host plants susceptible to the larvae resumed feeding and developed further to maturity. Second instar larvae were also able to establish new galls even though these galls were not as well developed as those caused by first instar larvae. Black currant plantations in northern Sweden were surveyed and local midge populations were found to be composed of either avirulent, virulent or a mixture of both midge types. Virulent midges were not restricted to plantations where resistant currant genotypes were grown. I conclude that, at least, two biotypes of the midge exist, and that those two are distinguished by the ability to gall and survive on `Storklas'.     

Interaction between relative humidity and resistance to Dasineura tetensi in blackcurrant

1 Infestation by the blackcurrant leaf midge Dasineura tetensi and galling incidence on a susceptible (cv. ‘Öjebyn’) and a moderately resistant (cv. ‘Storklas’) blackcurrant genotype was studied in the field for two midge generations in the same growing season. On the resistant genotype gall initiation is delayed. 2 The relation between infestation and galling incidence showed considerable variation between the two generations. Galling symptoms produced by the second midge generation were weak on the resistant as compared with the susceptible blackcurrant. 3 Because larval development of that generation coincided with a period of dry and warm weather, it is possible that larvae on the resistant genotype suffered desiccation to a greater extent than larvae on the susceptible genotype where gall development was stronger. 4 The possible interaction of relative humidity and expression of resistance was investigated in controlled environment experiments. The resistant ‘Storklas’ and a susceptible (‘7801–31’) currant genotype were studied at two constant relative humidities, 30 and 70%. 5 Larvae of D. tetensi suffered from higher mortality and reduced growth rate on both genotypes in the low humidity environment. 6 There was also a significant plant genotype by humidity interaction on larval performance; no galls were produced and no larvae completed development on the resistant currant at low humidity. 7 There was a trend for a positive correlation between larval length and larval density on a plant at low humidity but not at high humidity.     

Oxycephalomyia, gen. nov., and life history strategy of O. styraci comb. nov. (Diptera: Cecidomyiidae) on Styrax japonicus (Styracaceae)

A new genus Oxycephalomyia is described to contain the gall midge that was previously known as Asteralobia styraci (Shinji). Oxycephalomyia styraci, comb. nov., produces leaf vein galls on Styrax japonicus (Styracaceae). The adult of O. styraci is redescribed, and its full‐grown larva and pupa are described for the first time. The annual life cycle of the gall midge in northern Kyushu was clarified; the first instars overwinter in the galls on the host plant. However, the galls of O. styraci mature much later in the season than those of other gall midges with a similar life history pattern, and the durations of second and third larval instars are remarkably short. Such a life history pattern is considered to have an adaptive significance in avoiding larval parasitism, particularly by early attackers. The number of host axillary buds as oviposition sites decreased in bearing years and increased in off years, but there was no sign of oviposition site shortage even in bearing years, probably due to the low population density of the gall midge. An unidentified lepidopteran that feeds on galled and ungalled host buds and a Torymus sp. that attacks pupae of O. styraci were recognized as mortality factors of the gall midge.     

Preference–performance relationship in the gall midge Rabdophaga rosaria: insights from a common‐garden experiment with nine willow clones

1. Oviposition preferences of herbivorous insects are predicted to match offspring performance on different host taxa or on conspecific host genotypes. In gall‐inducing insects, host‐plant properties such as growth rate and gall size, which are determined by plant genotype and growing conditions, may have a significant impact on offspring performance and, hence, should influence oviposition site selection. 2. The present study investigated host preference of the European rosette willow gall midge Rabdophaga rosaria (Loew) in relation to offspring success on seven clones of Salix myrsinifolia Salisb. and two naturally hybridised S. myrsinifolia × phylicifolia L. clones growing in a replicated design in an experimental field under two fertilisation regimes. For each clone, the average growth rate, number of shoot tips, and leaf and gall size were determined, and their effects on midge preference and larval survival were examined. 3. Main shoot height, number of shoot tips, and gall size were significantly affected by clone. The midges clearly preferred certain clones over the others, but preferences were not related to willow growth traits or to gall size. Survival probability was higher in large than in small galls, but females did not prefer large‐leaved clones that produced the biggest rosette galls. Midge oviposition was also uncorrelated with prior rates of leaf‐rust infection and with feeding preferences of voles and folivorous insects. 4. The weak preference–performance relationship of R. rosaria within S. myrsinifolia is probably explained by evolutionary constraints that prevent generalist insects from achieving an ability to discriminate among conspecific hosts of variable quality.     

Rapid gall midge adaptation to a resistant willow genotype

Abstract 1 We conducted two experiments to investigate why a basket willow Salix viminalis L. genotype, known to be highly resistant to the leaf-roller gall midge Dasineura marginemtorquens (Bremi), should support very high gall densities in a field plantation at Tälle, south Sweden.
2 The first experiment was a field test of the hypothesis of fine-scale host adaptation in the gall midge/willow system. Support for the hypothesis would be established if midges originating from resistant willows and those originating from nearby susceptible willows differed in their abilities to initiate galls and complete development on resistant plants.
3 The objective of the second experiment was to explore whether there was a genetic basis to the trait for virulence in the midge population and to investigate any potential trade-offs this trait may entail.
4 Our results indicate that there was a fine-scaled microgeographic genetic structure to the midge population at Tälle. Midges originating from resistant plants had a heritable trait that enabled them to establish galls on resistant plants.
5 Midges able to initiate galls on the resistant genotype had longer developmental time on the susceptible genotype. This suggests that there is a physiological cost associated with being adapted to the resistant willow genotype.
6 We suggest that driving forces behind the observed host adaptation are selection imposed on the midge population by very strong willow resistance and restricted gene flow in the midge populations due to the special life history features of D. marginemtorquens .     

Life history strategy and taxonomic position of gall midges (Diptera: Cecidomyiidae) inducing leaf galls on Styrax japonicus (Styracaceae)

Four gall midge species (Diptera: Cecidomyiidae) that induce leaf galls on Styrax japonicus (Styracaceae) were identified to generic level based on larval morphology. Three of these gall midges, which induce whitish hemiglobular galls, flattened subglobular galls, and purple globular galls, respectively, were identified as three genetically distinct species of Contarinia, and the remaining species, which induces globular galls with dense whitish hairs, was identified as a species of Dasineura. Field surveys in Fukuoka, Japan, revealed that adults of these gall midges emerged and oviposited in late March to mid‐April at Mount Tachibana (approximately 200 m a.s.l.) and in late April to early May at Mount Sefuri (about 1050 m a.s.l.), coinciding with the leaf‐opening season of S. japonicus. Larvae of these gall midges mostly developed into third instars by June and then left their galls and dropped to the ground. These species therefore have a life history strategy that differs from that of another S. japonicus‐associated gall midge, Oxycephalomyia styraci, which overwinters as the first instar in ovate swellings, matures rapidly in spring, and emerges directly from the galls.     

Phytohormones in Japanese Mugwort Gall Induction by a Gall-Inducing Gall Midge

A variety of insect species induce galls on host plants. Liquid chromatographic/tandem mass spectrometric analyses showed that a gall midge (Rhopalomyia yomogicola) that induces galls on Artemisia princeps contained high levels of indole-3-acetic acid and cytokinins. The gall midge larvae also synthesized indole-3-acetic acid from tryptophan. Close observation of gall tissue sections indicated that the larval chamber was surrounded by layers of cells having secondary cell walls with extensive lignin deposition, except for the part of the gall that constituted the feeding nutritive tissue which was composed of small cells negatively stained for lignin. The differences between these two types of tissue were confirmed by an expression analysis of the genes involved in the synthesis of the secondary cell wall. Phytohormones may have functioned in maintaining the feeding part of the gall as fresh nutritive tissue. Together with the results in our previous study, those presented here suggest the importance of phytohormones in gall induction.     

Effect of sorghum resistant to Stenodiplosis sorghicola (Coquillett) (Diptera: Cecidomyiidae) on oviposition and development of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract Helicoverpa armigera oviposition preference for, and larval development on sorghum hybrids with differing resistance to sorghum midge, Stenodiplosis sorghicola , were investigated. When H. armigera larvae were fed seed of resistant and susceptible hybrids in the laboratory there were no differences in larval and pupal sizes or the rate of development. The same result was recorded when larvae fed on panicles on plants in a glasshouse. On some sampling occasions, significantly more eggs were laid on panicles of resistant hybrids in the field. This occurred when plants were in plots and also in a mixed planting. Midge-resistance status did not affect levels of egg parasitism. In a field study using recombinant inbred lines between a midge-resistant and a midge-susceptible line, no relationship was found between level of resistance and oviposition of H. armigera . We conclude that, although midge-resistant hybrids are sometimes preferred for oviposition by H. armigera, the resistance per se does not determine this preference. Egg survival, larval survival, development and resultant damage are not significantly affected by the midge-resistance status of the host.     

A rapid hypersensitive response associated with resistance in the willow Salix viminalis against the gall midge Dasineura marginemtorquens

Great genotypic variation in resistance against the gall midge Dasineura marginemtorquens Bremi (Diptera: Cecidomyiidae) exists within its willow host Salix viminalis L. (Salicaceae). In some resistant genotypes larvae die within 40 h after attempting to initiate galls. The present study tested the hypothesis that the hypersensitive response (HR) is involved in incompatible interactions between D. marginemtorquens and S. viminalis. By means of UV/blue light and visible light microscopy, we verified a rapidly (within 12 h after egg hatch) spreading cell death of an HR-type due to larval attack in resistant willow genotypes. Twelve h after egg hatch, the cell death had spread up to six cell layers in resistant S. viminalis genotypes whereas in susceptible genotypes only up to two cell layers were affected. In the groups of dead cells on the resistant genotypes accumulation of phenolics was observed within 24 h after egg hatch. The rapidity of the cell death induction, the early local accumulation of phenolic compounds, and the strong association of the cell death with larval mortality suggest that the formation of dead cells in resistant willow genotypes should be interpreted as a true HR.     

Life history strategy and adult and larval behavior of Macrodiplosis selenis (Diptera: Cecidomyiidae), a species that induces leaf‐margin fold galls on deciduous Quercus (Fagaceae)

Life historical, behavioral and ecological traits of Macrodiplosis selenis, which induces leaf‐margin fold galls on Quercus serrata, Q. mongolica and Q. dentata (Fagaceae) in Japan and South Korea, were studied. Daily activity and larval development indicate that M. selenis is a diurnal and univoltine gall midge. In April, females lay their eggs both on upper and under surfaces of fresh leaves. The duration of the egg stage varies from 5 to 9 days, depending on daily temperatures. Hatched larvae crawl to the upper surface of the leaf margin, where they start to induce galls. Larvae become full‐grown in October, drop to the ground in November and overwinter in cocoons on the ground, while larvae of congeners mature in May and drop to the ground in June. A relatively long period of the second larval stadium from July to October on the host trees seems to be effective for M. selenis in avoiding summer mortalities caused by predation and aridity on the ground and by ectoparasitoids that attack mature larvae or pupae on the host leaves. The spatial distribution pattern of M. selenis leaf galls is contagious and the mean gall density per leaf is significantly correlated with the mean crowding. This study adds new insights of life history strategy and adult and larval behavioral pattern to the ecological knowledge of gall midges, and these kinds of information are essential for further studies of M. selenis population dynamics and interactions with other Quercus‐associated herbivores.     

Asian chestnut gall wasp in Tuscany: gall characteristics,egg distribution and chestnut cultivar susceptibility

• 1 Preliminary investigations were carried out on Dryocosmus kuriphilus Yasumatsu on Castanea sativa Miller in Tuscany to assess variations in gall characteristics in coppice and high forest at two crown heights (height < 2 or 2–6 m), influence of bud size and bud position on oviposition rates and susceptibility of three cultivars.
• 2 Gall size may depend on various factors, including wasp population density. In the present study area, small galls (with one or two cells) were the most numerous in 2008, whereas larger galls (with more than three cells) prevailed in 2009.
• 3 Dryocosmus kuriphilus oviposition occurrence was influenced by both bud size and bud position. Buds with eggs tended to be larger in size compared with bud without eggs, suggesting that D. kuriphilus females prefer to lay eggs in larger buds (approximately 6 mm³) compared with smaller buds (approximately 3 mm³). The mean number of eggs per bud tended to decrease from the apical bud toward the basal bud.
• 4 Three C. sativa cultivars, Carpinese, Fusca and Cesurone, were examined. Fusca grafts had significantly more galls compared with Carpinese and Cesurone, whereas Cesurone grafts had more larvae per bud compared with Carpinese and Fusca. Overall, the Carpinese cultivar may be less susceptible to D. kuriphilus galling compared with the Fusca and Cesurone cultivars.

     

Internal structure of developing aucuba fruit as a defence increasing oviposition costs of its gall midges Asphondylia aucubae

Abstract. 1. Damage to juvenile plant tissues can cause reductions in fitness. Therefore, plants are expected to have evolved various defences for juvenile organs; however, so far, little attention has been paid to mechanical defence, as they have been considered to constrain the growth of juvenile organs. This study revealed that the dioecious tree Aucuba japonica uses mechanical defences to protect young developing fruit from the gall midge Asphondylia aucubae. 2. Young fruit of A. japonica have a hard layer of endocarp covering the integument. Midges oviposit on the surface of the integument, where larval chambers are later formed. The endocarp gradually becomes cracked as the embryo sac develops. 3. Oviposition by the midges is successful only when the ovipositors happen to pass through cracks in the endocarp. Thus, to successfully lay eggs, midges must insert the ovipositor repeatedly. This should decrease the fecundity of the midges, and subsequently their infectiousness, because their adult lifespan is short and they do not consume food during this time. 4. Expansion of the cracks in the endocarp simplifies oviposition over time; however, the embryo sac continues to grow, increasing its volume relative to that of the ovule. This appears to deplete available space and tissue used in the construction of larval chambers, gradually making the fruit less susceptible to midge attacks. 5. The temporary nature of this defence should prevent it from constricting the growth of young fruit. This exemplifies a novel strategy for a mechanical defence of young developing plant tissues.     

Oviposition by autumn gum moth (Mnesampela privata) in relation to Eucalyptus globulus defoliation, larval performance and natural enemies

Abstract 1 Eucalyptus globulus Labill. exhibited consistent intraspecific variation in oviposition choice by Mnesampela privata (Guenée) (Lepidoptera: Geometridae) in field surveys of host plants that had been designated, based on the prior season's defoliation levels, as resistant or susceptible to M. privata. At both field sites significantly fewer egg batches were found on resistant compared to susceptible trees. 2 In cage bioassays we demonstrated a significant oviposition preference by M. privata, with over two‐fold greater number of egg batches laid on foliage sprigs from susceptible compared to resistant trees. Despite differences in tree oviposition choice, we found no evidence in the field or the laboratory of adult females adjusting egg batch size in accordance to oviposition preference. 3 Caged larval survival, development times and pupal weight did not vary significantly between resistant and susceptible trees. Similarly, in a laboratory feeding experiment, neonates utilized resistant and susceptible foliage equally. 4 Larval mortality in the field attributed to natural enemies did not vary significantly between resistant and susceptible trees, nor did the percentage of Telenomus sp. parasitism of M. privata eggs within a batch and batches per tree. 5 Failure to associate either larval performance or natural enemy efficacy with the observed intraspecific variation in E. globulus susceptibility to M. privata oviposition indicates that some other unidentified factors drive the evolution of host selection for oviposition.     

High oviposition rate ofDasineura marginemtorquens onSalix viminalis genotypes unsuitable for offspring survival

A short history of contact betweenDasineura marginemtorquens and resistant genotypes ofSalix viminalis has previously been suggested as an explanation for a poor correspondence between female oviposition preference and larval performance on certain host types. The discovery of resistant wild willows instigated a quest for an alternative hypothesis. This study presents data from laboratory and field experiments that were designed to detect variability in host preferences of individual members of the midge population. Field tests showed that the oviposition site choices of females followed the same random-natured pattern as choices of females in laboratory situations. Ovipositing females did not distinguish between willow genotypes with very high or very low larval mortality. No differences in post-alighting behavior of midges on the two willow types were discovered. Observations revealed that females usually remained and oviposited on the first plant with which they came into contact, regardless of genotype.     

Intra‐plant heterogeneity influences the preference and performance of juveniles and adults of a gall midge

Abstract 1. Field studies were conducted to evaluate the preference and performance of a gall‐inducing midge (Harmandia tremulae) within the crown of trembling aspen (Populus tremuloides). Females did not select oviposition sites preferentially within leaves, but did lay preferentially on young leaves. 2. Larvae were the only life stage involved in gall site selection within leaves and in gall initiation and development. Gall size, which was positively related to survival, was highest for galls on mid veins that were located close to the petiole. However, one‐third of galls were located on lateral veins and most galls were not adjacent to the petiole, indicating that many larvae choose sub‐optimal gall initiation sites. 3. Gall density was positively associated with leaf length, and leaf length, was positively associated with gall size. However, gall density per leaf was not related to larval survival in galls. This latter result may be a result of an observed inverse relationship between gall size and gall density for similar‐sized leaves. 4. The results partially support the plant vigour and optimal plant module size hypotheses, which predict that galler fitness in successfully induced galls should be highest on large, fast‐growing plant modules. The lack of a strong preference‐performance link supports the confusion hypothesis, which predicts that oviposition and gall site selection may often be suboptimal in systems where galler lifespan is short. This study suggests that small‐scale variations in plant quality within leaves, can render gall site selection by juveniles as important as that previously reported for adult females.     

Intraspecific variation in beech scale populations and in susceptibility of their host Fagus sylvatica

ABSTRACT.

• 1 Within a seed orchard in southern England, beech trees (Fagus sylvatica L.) belonging to three clones were artificially infested by introducing beech scale larvae (Cryptococcus fagisuga Lind. Homoptera: Coccidae) into small cages attached to the bark.
• 2 Some larvae developed to fecund adults on trees of two susceptible clones but all failed to develop on a third, resistant clone.
• 3 Within susceptible clones, survival of larvae on individual trees was positively related to their degree of natural infestation.
• 4 Larvae deriving from several separate trees differed significantly in their ability to survive when inoculated onto trees of susceptible clones.
• 5 Five forest trees which acted as both donors of larvae and as hosts for artificial inoculation were each inoculated with larvae from all five trees.
• 6 There was significant variation in survival of inoculated larvae both between the host trees and between sources of larvae on each host.
• 7 Survival of larvae reinoculated onto their original host was significantly higher than that of larvae originating from other trees.
• 8 Fecundity of adults on the forest trees was positively correlated with the probability of inoculated larvae surviving to the adult stage.

     

Influence of plant genotype and environment on oviposition preference and offspring survival in a gallmaking herbivore

Summary Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.     

Oviposition and larval survival of Dasineura tetensi on four blackcurrant Ribes cultivars

Monitoring of an unsprayed infested fieldsite using watertraps in S.E. Kent revealed four generations of Dasineura tetensi (Rubs) (Diptera: Cecidomyiidae) occurring between April and September 1996. Ribes nigrum L. cultivars 'Baldwin' (susceptible), 'Ben Alder' (susceptible) and 'Ben Connan' (resistant) were sampled for eggs in the field and assessed for midge damage throughout the season. Oviposition was indiscriminate, but plant damage varied significantly between cultivars. In laboratory choice experiments, mated female midges showed no preference between susceptible shoots of 'Ben Alder' and resistant shoots of 'Ben Connan' for oviposition. Olfactory responses of D. tetensi to leaf volatiles of 'Ben Alder' and 'Ben Connan' were also tested in a 4-way olfactometer. Mated females did not discriminate between volatiles of susceptible and resistant host plants. Larvae reared on cv. 'Ben Connan' shoots were significantly smaller than those reared on shoots of cv. 'Ben Alder'. Larval antibiosis and not female antixenosis appears to be the main mechanism for resistance to D. tetensi in 'Ben Connan'.     

Timing of growth determines fitness and performance of a galling insect on willow

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