Low genetic variability in a Utah cherry-infesting population of the apple maggot, Rhagoletis pomonella

The apple maggot fly, Rhagoletis pomonella (Walsh), has only recently been found in Utah infesting sour cherry, Prunus cerasus L. An electrophoretic comparison of flies from Utah cherries with flies from Illinois hawthorns, Crataegus mollis (T. & G.) Scheele (a native host within the native range of the fly), show a marked reduction of genetic variability in the Utah sample. This result is indicative of a genetic bottleneck associated with the establishment of the apple maggot population in Utah cherries.

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Résumé R. pomonella (Walsh), est originaire de Crataegus dans l'Amérique du N.E. Il attaque de nombreux autres fruits, y compris les pommes et les cerises aigres (Prunus cerasus). La mouche a été récemment signalée en Utah, à la fois sur cerises et sur Crataegus douglasii. Nous avons comparé les niveaux de variabilité génétique d'une population de l'Utah contaminant les cerises et d'une population de l'Illinois contaminant C. mollis (la population de l'Illinois est représentative des niveaux de variabilité génétique dans l'aire d'origine de la mouche).La variabilité génétique à 17 loci a été évaluée par électrophorèse sur gel d'amidon. 10 de ces loci sont polymorphes dans la population d'Illinois, mais seulement 4 dans la population de l'Utah. Les fréquences alléliques de ces 4 loci de R. pomonella diffèrent significativement en Utah et en Illinois. La population de l'Utah présente nettement moins d'allèles par locus, un plus faible pourcentage de loci polymorphes et une hétérozygotie moyenne plus faible que la population de l'Illinois. Tous ces résultats sont conformes aux conséquences prévisibles d'un goulot d'étranglement.Deux explications existent pour cette perte de variabilité, toutes les deux liées à la combinaison de la faible taille de la population et de la dérive génétique ultérieure. Pour la première, la colonisation du cerisier par les mouches venant de Crataegus peut avoir provoqué un goulot d'étranglement génétique. Au contraire, la réduction de la variabilité peut avoir été la conséquence de la colonisation de l'Utah par R. pomonella. Nous retenons cette dernière comme la cause la plus vraisemblable de la variabilité génétique de la population de R. pomonella contaminant les cerises de l'Utah.

     

Influence of protein hydrolysate baits on oviposition in the apple maggot, Rhagoletis pomonella

Ovipositional responses of apple maggot (AM), Rhagoletis pomonella (Walsh), females were studied in the laboratory on apples (var: Golden Delicious) treated with different rates of four protein hydrolysate baits in choice and no-choice tests. Protein hydrolysate baits at rates of 0.5 and 1% had no significant effect, but oviposition was greatly reduced at higher rates of 5 and 10%. Apple maggot females exposed to apples treated with protein hydrolysate baits at a rate of 10% made 41–71% fewer punctures and laid 41–73% fewer eggs than in untreated control. No oviposition activity was shown on apples treated with 25 and 100% Nulure®. In no-choice tests the AM females laid 75–96% fewer eggs in apples treated with 10 and 25% Nulure compared to controls and no oviposition occurred in apples treated with 100% Nulure. Apple maggot females arrived in similar numbers on apples treated with 10% Nulure and untreated apples, but only 5% of those arriving on Nulure-treated apples showed ovipositor boring with no egg deposition while 60% of females arriving on untreated apples showed ovipositor boring activity and laid an average of 2.5 eggs per apple. In another experiment, individual AM females displayed similar behavioral responses to 10% Nulure-treated apples; none of the 56 females tested on treated apples displayed ovipositor boring activity, but 59% of the females (N=56) tested on untreated apples displayed ovipositor boring within 5 min of their arrival. Ninetyeight percent of AM females stayed and fed on fruit surfaces for 5 min on Nulure-treated apples without ovipositor boring compared to only 2% on untreated apples. Of the females that arrived on untreated apples, 39% flew away within 5 min without ovipositor boring compared to only 2% of those that arrived on Nulure-treated apples. Results of these two behavioral experiments suggest that upon arrival on a protein bait-treated apple, an apparent change of behavior occurs in AM females and instead of attempting to oviposit, they attempt to feed on fruit surfaces resulting in reduced oviposition activity. These results indicate that the feeding and oviposition-related activities of AM females are probably mutually exclusive and that the feeding behavior preempts oviposition activities on host fruits treated with higher rates of protein hydrolysate baits.     

Prior experience influences the fruit residence of male apple maggot flies,Rhagoletis pomonella

Male apple maggot flies spend considerable time residing on individual host fruit as territories on which they force-copulate arriving females in search of oviposition sites. Here, we present evidence from investigations in nature and the laboratory that shows the propensity of males to reside on a hawthorn or apple fruit as a territory is significantly modifiable through prior experience with fruit and, hence, involves learning. Previous studies revealed that after a female apple maggot fly, Rhagoletis pomonella, arrived on a host hawthorn or apple fruit, its propensity to accept or reject that fruit for egg-laying was similarly modifiable through prior fruit-exposure experience and also involved learning. We discuss how host fruit learning in males and females, in concert with genetic-based differences in host fruit residence and acceptance behavior between populations of flies originating from hawthorn and apple, could give rise to a reduction in gene flow between populations of flies on these two host types.     

Attracticidal spheres for controlling apple maggot flies: commercial-orchard trials

Wooden pesticide-treated red spheres and biodegradable sugar/flour pesticide-treated red spheres were compared with wooden sticky-coated red spheres and insecticide sprays for controlling apple maggot flies, Rhagoletis pomonella(Walsh) (Diptera: Tephritidae), in small blocks of apple trees in Massachusetts commercial orchards in 1997, 1998 and 1999. Pesticide-treated spheres received a coating of 70% latex paint, 20% feeding stimulant (sucrose), and 10% formulated insecticide (containing 20% imidacloprid). To replenish sucrose lost during rainfall, wooden spheres were capped with a disc comprised of hardened sucrose that seeped onto the sphere surface, whereas the surface of sugar/flour spheres received sucrose that seeped from the interior. Each year, each of the 24 perimeter trees of each non-sprayed block received a sphere baited with butyl hexanoate (an attractive component of host fruit odor), with the intent of intercepting immigrating flies. Based on captures of flies on unbaited sticky-coated red spheres placed near the center of each block and on periodic samples of fruit for injury, there was a consistent pattern of treatment performance. Each year, sticky-coated spheres were only slightly less effective than two or three sprays of organophosphate insecticide, sugar/flour pesticide-treated spheres were only slightly less effective than sticky spheres, and wooden pesticide-treated spheres were least effective. Versions of pesticide-treated spheres used in 1999 were more durable than those used in previous years, but further improvement is needed before either wooden or sugar/flour pesticide-treated spheres can be recommended for grower use.     

Learning of apple fruit biotypes by apple maggot flies

Previously, we showed that after a female apple maggot fly, Rhagoletis pomonella,arrives on a host hawthorn or apple fruit, its propensity to accept (bore into) or reject that fruit prior to egg deposition can be modified by previous ovipositional experience with one or the other species and, hence, involves learning. Here, we present both field and laboratory evidence indicating that females also are able to learn characteristics of three different apple biotypes or cultivars: Early Macintosh, Red Delicious, and Golden Delicious. We suspect that females learn to discriminate among these three cultivars on the basis of differences in chemical stimuli among cultivars. The effect of fruit cultivar learning was not as strong as the effect of fruit species learning.     

Genetic variation among apple and hawthorn host races of Rhagoletis pomonella across an ecological transition zone in the Mid-Western United States

In previous studies, we have shown that apple and hawthorn populations of Rhagoletis pomonella (Diptera: Tephritidae) represent partially reproductively isolated and genetically differentiated host races; a result consistent with predictions of sympatric speciation models. The geographic pattern of allozyme variation for these flies is complex, however, as inter-host differences are superimposed on latitudinal allele frequency clines within the races. In addition, pronounced allele frequency shifts exist among R. pomonella populations across three major ecological transition zones in the mid-western United States. This suggests that selection related to environmental heterogeneity is responsible for the allele frequency shifts, but does not rule out secondary contact as an alternative possibility. Resolution of this issue is important, because if secondary contact is involved, then we would have to reassess the relationship host race formation has with speciation in the R. pomonella group.Here, we present results from a detailed genetic analysis of fly populations spanning the deciduous/prairie transition zone near the border between the states of Wisconsin and Illinois. Allele frequencies for hawthorn populations within the zone formed spikes, rather than the expected steps, and these frequency peaks correlated with variation in local ambient temperature conditions. Ambient temperature, and not secondary contact, therefore appears to be an important determinant of the shape of R. pomonella allele frequency clines. Allele frequency heterogeneity was also observed among apple populations, but was less pronounced compared to that for hawthorn flies. This suggests that ambient temperature differentially affects the host races, possibly through differences in the fruiting phenologies of apple and hawthorn trees. Several pairs of linked loci displayed concordant allele frequency changes and were in disequilibrium among both apple and hawthorn populations along the Wisconsin/Illinois transect. Although we do not know the reason for the observed pattern of disequilibrium, site to site variation in levels of inter-host migration, coupled with selection, seem the most likely explanations. We conclude by discussing how host specific adaptations, such as those associated with ambient temperature, may interact with host recognition traits to drive the sympatric speciation process for R. pomonella group flies.     

Multiple Mating and Reproductive Success of Male and Female Apple Maggot Flies, Rhagoletis pomonella (Diptera: Tephritidae)

Mature male and female apple maggot flies mated frequently on a field-caged host tree during a 14-day study. Each sex averaged one mating per day (mean of 1.0 ± 0.1), but some females mated up to eight times per day and some males up to six times per day. Reproductive success was estimated based both on observed numbers of matings in the field cage and on previous work relating fecundity and fertility to female mating status. Male and female flies did not differ in mean or variance in reproductive success, indicating that this is a polygamous mating system consisting of both polygyny and polyandry. We discuss the significance of this with regard to the framework of insect mating control and frequency proposed by others. We also discuss behaviors (such as movements, agonistic encounters, occurrences on fruit, and ovipositions) that do and do not show correlations with mating success in apple maggot flies.     

The population genetics of the apple maggot fly,Rhagoletis pomonella and the snowberry maggot,R. zephyria: implications for models of sympatric speciation

True fruit flies belonging to the Rhagoletis pomonella (Diptera:Tephritidae) sibling species complex possess several attributes consistent with a history of sympatric divergence via host plant shifts. Here, we investigate whether hybridization and genetic introgression is occurring between two members of the group, Rhagoletis pomonella (Walsh), whose primary hosts are domestic apples (Malus pumila) and hawthorns (Crataegus spp., and R. zephyria (Snow) whose host is snowberries (Symphoricarpos spp.). These flies are important because they appear to be at a transition stage between taxa reproductively isolated solely on the basis of host plant-related adaptations and those that have evolved additional non-host dependent sterility and inviability. Observing extensive hybridization and introgression between R. pomonella and R. zephyria would have major repercussions for current models of sympatric speciation.In a survey of allozyme and mitochondrial DNA (mtDNA) variation for 1105 flies collected throughout the northern United States we found two results suggesting that low level hybridization occurs between R. pomonella and R. zephyria. (1) Two flies reared from snowberries and one fly reared from hawthoms had genotypes indicative of them being R. pomonella and R. zephyria, respectively. Rhagoletis pomonella and R. zephyria adults may therefore occasionally frequent each others host plant, providing the opportunity for hybridization. (2) Four flies collected from hawthorns and one from snowberries had genotypes that made them likely to be F1 hybrids.Likelihood analysis revealed the data were also consistent with an hypothesis of shared ancestral alleles (i.e., the pattern of genetic variation could also be explained by R. pomonella and R. zephyria sharing alleles/haplotypes whose origins date to a common ancestor). We estimated that, in the absence of interspecific mating, random assortment of genes within R. pomonella and R. zephyria populations would produce an average of 5.4 flies with genotypes suggesting they were F1 hybrids – a number equivalent to the 5 putative F1 hybrids observed in the study. Our results therefore underscore the difficulty in distinguishing between hypotheses of low level introgression and shared ancestral polymorphism. But even if hybridization is occurring, the data suggest that it is happening at a very low and probably evolutionarily insignificant level (perhaps 0.09% per generation), consistent with sympatric speciation theory. Future tests are discussed that could help resolve the hybridization issue for R. pomonella and R. zephyria.     

Isolation and characterization of microsatellite loci from the apple maggot fly Rhagoletis pomonella (Diptera: Tephritidae)

We report the isolation and development of 81 novel primers for amplifying microsatellite loci in the Rhagoletis pomonella sibling species complex, and the sequencing, characterization and analysis of basic population genetic parameters for nine of these genes. We also report the successful cross‐species amplification of several of these loci. The R. pomonella sibling species complex is a textbook example of genetic differentiation in sympatry via host‐plant shifting. Microsatellite markers can be useful for mapping host‐plant‐associated adaptations in Rhagoletis that generate reproductive isolation and facilitate speciation, as well as for resolving the genetic structure and evolutionary history of fly populations.     

Prior experience affects the visual ability ofRhagoletis pomonella flies (Diptera: Tephritidae) to find host fruit

Females of the apple maggot fly,Rhagoletis pomonella, were allowed for 3 days to alight upon and oviposit in green or red 18- to 20-mm hawthorn host fruit (Crategus mollis) or green or red 45- to 55-mm apple host fruit (Malus pumila) hung from branches of potted host trees in field enclosures. Subsequently, when females were released individually on potted host trees harboring fruit of one of these types, their ability to find fruit of unfamiliar size proved unaffected by prior experience with fruit but their ability to find fruit of unfamiliar color was significantly affected. Specifically, females exposed to red hawthorns or red apples were less able to find green hawthorns or green apples than were females experienced with either of the latter fruit types. Fruit odor was found to have no effect on female ability to find familiar compared with unfamiliar green fruit. In contrast, a difference in size (or surface chemistry) between familiar and unfamiliar fruit but not a difference in fruit color had a significant negative influence on the propensity of alighting females to bore into unfamiliar fruit. Three bouts of experience with alighting upon and ovipositing into fruit over a period of about 1 h had no detectable effect on female ability to find unfamiliar fruit but did reduce propensity to bore into unfamiliar fruit. Our findings are discussed in relation to insect ability to learn visual and chemical stimuli of resources and insect propensity to form host races. We also discuss the potential impact of our findings on nonpesticidal, behavioral methods of managingR. pomonella in commercial apple orchards.     

The effects of climate, host plant phenology and host fidelity on the genetics of apple and hawthorn infesting races ofRhagoletis pomonella

True fruit flies belonging to theRhagoletis pomonella (Walsh) sibling species complex have been proposed to speciate sympatrically by shifting and adapting to new host plants. Here, we report the results from a series of ecological and genetic experiments conducted at a study site near Grant, Michigan, U.S.A., aimed at clarifying the relationship between host specialization and reproductive isolation for these flies. Our findings indicate that apple (Malus pumila) and hawthorn (Crataegus mollis) infesting populations ofR. pomonella are partially allochronically isolated. Differences in the timing of adult eclosion account for part of the allochronic divergence, as apple adults emerge approximately ten days earlier than hawthorn flies in the field. Genetic analyses across different life history stages of the fly show that adults do not randomly move between apple and hawthorn trees, but trend to attack the same species of plant that they infested as larvae. Estimates of interhost migration from the allozyme data suggest that from 2.8 to 10% of the apple population is of hawthorn origin and that over 20% of the hawthorn population is of apple origin. The length and quality of the growing season appear to affect the genetic composition of the host races, as allele frequencies in the hawthorn population are correlated with ambient temperature and rainfall during the spring of the preceding year. Finally, allele frequencies for six allozyme loci displaying host associated differentiation also show significant linear regressions with the timing of adult eclosion within both races. These regressions establish a link between allozyme loci displaying inter-host differentiation and a developmental trait (adult eclosion) responsible for partially isolating the races. The slopes of the regressions are paradoxical, however, as they suggest that apple adults should eclose later, not earlier, than hawthorn flies. We conclude by discussing potential resolutions to the eclosion time paradox.     

Apple maggot fly response to red sphere traps in relation to fly age and experience

Responses of the apple maggot fly,Rhagoletis pomonella (Walsh), to 8 cm red sticky sphere traps, baited with the synthetic fruit odor butyl hexanoate, were investigated in field-caged apple trees containing green or red Gravenstein apples. Trap capture rate and the probability of oviposition in apples before capture generally increased with female age and number of mature eggs in ovaries. Two days of pre-test exposure of mature females to red hawthorns, green Red Delicious apples or green Gravenstein apples had no significant effect on the likelihood of a fly finding a red sphere. However, before capture on a sphere or departing a tree, hawthorn-exposed females found significantly fewer apples and laid significantly fewer eggs than females exposed to Gravenstein apples. Variation in duration of pre-test exposure (1–4 days) of flies to Gravenstein apples had no detectable influence on female response to apples or to a red sphere in a test tree. The relevance of these findings to effectiveness of sphere traps, forR. pomonella control in commercial orchards is discussed.     

Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot fly,Rhagoletis pomonella

Evidence suggests that the apple maggot, Rhagoletis pomonella (Diptera: Tephritidae) is undergoing sympatric speciation (i.e., divergence without geographic isolation) in the process of shifting and adapting to a new host plant. Prior to the introduction of cultivated apples (Malus pumila) in North America, R. pomonella infested the fruit of native hawthorns (Crataegus spp.). However, sometime in the mid-1800s the fly formed a sympatric race on apple. The recently derived apple-infesting race shows consistent allele frequency differences from the hawthorn host race for six allozyme loci mapping to three different chromosomes. Alleles at all six of these allozymes correlate with the timing of adult eclosion, an event dependent on the duration of the overwintering pupal diapause. This timing difference differentially adapts the univoltine fly races to an approximately 3- to 4-week difference in the peak fruiting times of apple and hawthorn trees, partially reproductively isolating the host races. Here, we report finding substantial gametic disequilibrium among allozyme and complementary DNA (cDNA) markers encompassing the three chromosomal regions differentiating apple and hawthorn flies. The regions of disequilibrium extend well beyond the previously characterized six allozyme loci, covering substantial portions of chromosomes 1, 2, and 3 (haploid n = 6 in R. pomonella). Moreover, significant recombination heterogeneity and variation in gene order were observed among single-pair crosses for each of the three genomic regions, implying the existence of inversion polymorphism. We therefore have evidence that genes affecting diapause traits involved in host race formation reside within large complexes of rearranged genes. We explore whether these genomic regions (inversions) constitute coadapted gene complexes and discuss the implications of our findings for sympatric speciation in Rhagoletis.     

Behavioral evidence for host races in Rhagoletis pomonella flies

Summary One of the most controversial putative cases of host race formation in insects is that of the apple maggot fly, Rhagoletis pomonella (Diptera: Tephritidae). A principal cause of the controversy is lack of relevant data. In laboratory and field enclosure experiments, we compared the host acceptance behavior of sympatric populations of flies originating from naturally infested hawthorn (the native host) and apple (an introduced host) in Amherst, Massachusetts or East Lansing, Michigan. In general, hawthorn fruit were accepted for ovipositional attempts nearly equally by apple and hawthorn origin females, whereas apples were accepted much more often by apple than hawthorn origin females. Similarly, males of apple and hawthorn origin exhibited about equal duration of residence on hawthorn fruits as sites at which to acquire potential mates, while males of apple origin tended to reside substantially longer than males of hawthorn origin on apples. Irrespective of fly origin, both sexes always responded more positively to hawthorn fruit than to apples. Because all flies assayed were naive (ruling out effects of prior host experience of adults) and because tests revealed no influence of pre-imaginal fruit exposure on pattern of host fruit acceptance by females, the combined evidence suggests the phenotypic differences we observed in host response pattern between hawthorn and apple origin flies may have an underlying genetic basis. Further tests showed that while larval progeny of flies of each origin survived better in naturally growing hawthorn fruit than in naturally growing apples, there was no differential effect of fly origin on larval survival ability in either host. We discuss our findings in relation to restriction in gene flow between sympatric populations of R. pomonella and in relation to current models of host shifts in insects.     

Visual responses of apple maggot flies, Rhagoletis pomonella (Diptera: Tephritidae): Orchard studies

Apple maggot flies, Rhagoletis pomonella, were found to be highly responsive to visual stimuli, with maximum responses dependent upon color, shape, and size characteristics of the stimulus. Each sex had a significant preference for yellow 30×40 cm rectangles over green, orange, red, blue, violet, black, white and clear rectangles of the same size. On the other hand, each sex had a significant preference for red, blue, violet, dark organge, and black spheres 7.5 cm in diameter over green, light orange, yellow, white, and clear spheres of the same size. Both sexes had a significant preference for 7.5 cm spheres over equivalent-sized cubes, cylinders and rectangles. As the diameter of a sphere was increased from 7.5 to 45 cm, there was an orderly and significant decrease in the attractiveness of those that were darkcolored (red) but a progressive and significant increase in the attractiveness of those that were yellow. The flies did not congregate on trees with apples or respond strongly to 7.5 cm, darkcolored spheres until they were sexually mature (at least in the case of females). Relative to other colors, females were often significantly more attracted than males to yellow. The attractiveness of an olfactory stimulus eliciting feeding-type reactions was enhanced to a substantial degree when employed in conjunction with a 30×40 cm yellow rectangle, but to only a very slight degree when in conjunction with a 7.5 cm red sphere.Evidence presented suggests the following explanation for these findings. The flies are attracted to large surfaces of yellow because they react to yellow, on the basis of true color discrimination, as if it were foliage on which to find food. On the other hand, they are attracted to small, dark-colored spheres because they react to such spheres as they react to apples, which are the oviposition site and which also appear to serve as a rendezvous site for the sexes in mating activity. Once the flies have arrived on apple trees, they detect apples solely through vision. Red, blue, violet, dark orange, and black small spheres are preferred over small spheres of other colors on the basis that they stand out in strongest contrast against the background and not on the basis of true color discrimination. Large, dark-colored spheres are unattractive because the range of sphere sizes eliciting positive responses is near the size of an apple.These findings have proven useful in designing effective devices for sampling orchard populations of the flies and may also prove useful in fly control.

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Zusammenfassung Imagines von Rhagoletis pomonella reagieren positiv auf visuelle Reize, wobei Farbe, Form und Grösse des Stimulus von ausschlaggebender Bedeutung sind. Beide Geschlechter zeigten in den Versuchen eine signifikante Präferenz für gelbe Rechtecke (30×40 cm), die sie häufiger anflogen als grüne, orange, rote, blaue, violette, schwarze, weisse und farblose Flächen gleicher Grösse. Ebenso zeigte es sich, dass Kugeln von 7.5 cm Durchmesser von den Fliegen eindeutig Kuben, Zylindern und Rechtecken gleicher Grössenordnung für den anflug bevorzugt wurden. Mit zunehmendem Durchmesser der Kugeln von 7.5 cm bis 45 cm konnte eine abnehmende Anziehung von dunkel gefärbten Kugeln, jedoch eine zunehmende Anziehung von gelben Kugeln auf die Fliegen beobachtet werden. Vor der Geschlechtsreife reagierten die Weibchen nicht in ausgeprägtem Maße auf die 7.5 cm dunkel gefärbte grossen Kugeln, sondern wurden mehr als die Männchen von Gelb angezogen. Die Wirkung eines Nahrungsköders wurde in Kombination mit einer gelben Fläche von 30×40 cm signifikant erhöht, während die Kombination des Köders mit einer roten Kugel von 7.5 cm Durchmesser die Anziehung von Weibchen nur unwesentlich erhöhte.Die Resultate lassen sich damit erklären, dass die Fliegen durch gelbe Flächen angezogen werden, weil sie dank Diskriminierung der Farbe auf Gelb reagieren als wäre es Blattwerk, auf dem sich die Nahrungsquellen befinden. Auf der anderen Seite werden sie von dunklen Kugeln in den Dimensionen eines Apfels angezogen, da im Freiland Äpfel Orte für die Eiablage und offenbar Treffpunkte für die Geschlechter darstellen. Sobald die Fliegen den Apfelbaum beflogen haben, orientieren sie sich allein mit dem Gesichtssinn gegen die Früchte. Kleinere rote, blaue, dunkel orange, violette und schwarze Kugeln werden anders gefärbten vorgezogen, weil sie den grössten Kontrast gegenüber dem Hintergrund bilden und nicht auf Grund echter Farbdiskriminierung. Grosse dunkle Kugeln verlieren ihre Wirkung auf die Fliegen, weil nur Kugeln in der Grössenordnung eines Apfels positive Reaktionen auslösen.Diese Befunde haben sich als nützlich erwiesen in der Ausarbeitung von Sammelmethoden für Freilandpopulationen und dürften auch in der Bekämpfung der Apfelfliegen Anwendung finden.

     

Effects of pH on feeding responses in the apple maggot fly,Rhagoletis pomonella (Diptera: Tephritidae)

Phagostimulatory effects of pH values of sucrose on Rhagoletis pomonella adults were studied in the laboratory. Flies were standardized for age, diet and food deprivation. Two presentation schemes were employed. The first varied pH value (3.0-10.0) with sucrose concentration kept constant at 40%. The second varied both sucrose concentration (8%, 24% and 40%) and pH value (5.0-8.0). Fly feeding propensity was evaluated by observation of fly acceptance or rejection of sucrose and duration of feeding. When tested on red wooden spheres treated with 40% sucrose, fly feeding acceptance was significantly greater when pH ranged from 5.0 to 8.0, and duration of feeding was significantly longer at pH 6.0-7.0. At pH /=8.0, feeding propensity was significantly reduced. Decrease in sucrose concentration significantly increased fly sensitivity to pH. Males were more responsive to varying pH than females. The sucrose pH shown to stimulate maximal feeding response was 6.4. Such information is relevant to formulation improvement of a coating mixture of sucrose and insecticide applied to red spheres as part of apple maggot fly control programs.     

Pleiotropy of insecticide resistance in the codling moth, Cydia pomonella

The codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), has developed resistance to various insecticides. Relative fitness of one susceptible strain (Sv) and two strains selected for resistance to diflubenzuron (Rt) and deltamethrin (Rv), respectively, was measured in the absence of insecticide selection pressure. Mating rate, fecundity, fertility, developmental time, fifth instar weight, and adult longevity were compared. Both resistant strains were less fecund and fertile, developed more slowly, weighed less, and had shorter life-spans than the susceptible strain. These results indicate that biological constraints are associated with insecticide resistance in the codling moth. We also found that fitness estimates of the Rv strain did not differ statistically from those of the Rt strain. Enhanced mixed-function oxidase and glutathione-S-transferase activities have been shown to be involved in insecticide resistance in both Rt and Rv strains. This suggests that the fitness cost described in both resistant strains was mainly associated to metabolic resistance. The impact of such deleterious pleiotropy of insecticide resistance in C. pomonella in terms of resistance management in the field is discussed.     

The hymenopteran parasitoids of light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) in Australia

Abstract  Epiphyas postvittana (Walker) is a serious pest of a number of horticultural crops including grapes in Australia and New Zealand. This study brings together information on the parasitoid complex associated with E. postvittana that previously was fragmented and largely inaccessible. We include species reared during a 3-year study of the parasitoids of E. postvittana in the vineyards of the Coonawarra region, South Australia, material from several Australian agricultural insect collections and records from the literature. An illustrated key is presented for 25 species of parasitoids and hyperparasitoids associated with E. postvittana , along with information on the taxonomy, identification, distribution and biology of each species. Taxa newly recorded from this host are Perilampus sp. (Perilampidae), and six species of Ichneumonidae: Euceros sp., Labium sp., Netelia sp., Plectochorus sp., Temalucha minuta (Morley) and Eriborus epiphyas sp. n., the latter species being described in full.     

The cost of reproduction in rosehip flies,Rhagoletis basiola: Eggs are time

Summary The hypothesis was tested that egg-laying by rosehip fruit flies,Rhagoletis basiola, would lead to reduced adult survivorship compared with oviposition-deprived flies, through experimental manipulation. Oviposition-deprived flies were assigned to two groups: mated and spinster. All three treatment groups were permitted to spend equal time and energy exploring and handling hosts but only the former group were allowed to deposit eggs (three/day). Results clearly showed that ovipositing flies suffered significantly higher mortality rates and that mated and spinster flies produced similar survivorship patterns. I consider these results both as a foraging problem (i.e. laying eggs in inferior hosts can cost both eggs and life) and as a more general life history problem.