Evaluation of trap deployment patterns for behavioural control of apple maggot flies (Dipt., Tephritidae)

Abstract: We evaluated three different deployment patterns of sticky red sphere traps, baited with a five-component blend of synthetic attractive fruit odour and placed on perimeter apple trees bordering adjacent habitat (front-row trees), for control of apple maggot flies, Rhagoletis pomonella (Walsh), in small plots of apple trees (about 30 × 30 m) in Massachusetts commercial apple orchards. Degree of fly penetration from front-row to interior apple trees was assessed for R. pomonella of wild origin and for marked adults released in habitats adjacent to front-row trees. Traps placed 10  and 5 m apart on front-row trees or grouped on a single central front-row tree performed as well as grower-applied insecticide sprays in preventing penetration of plots by wild and released flies and in preventing fruit injury . This was equally true for plots whose front-row trees consisted of cultivars comparatively susceptible to apple maggot as for plots whose front-row trees were comprised of comparatively tolerant cultivars. It was also true for each seasonal period during which sampling for treatment performance occurred.     

Intra- and interspecific competition and host race formation in the apple maggot fly,Rhagoletis pomonella (Diptera: Tephritidae)

Intra- and interspecific resource competition are potentially important factors affecting host plant use by phytophagous insects. In particular, escape from competitors could mediate a successful host shift by compensating for decreased feeding performance on a new plant. Here, we examine the question of host plant-dependent competition for apple (Malus pumila)- and hawthorn (Crataegus mollis)-infesting larvae of the apple maggot fly, Rhagoletis pomonella (Diptera: Tephritidae) at a field site near Grant, Michigan, USA. Interspecific competition from tortricid (Cydia pomonella, Grapholita prunivora, and Grapholita packardi) and agonoxenid (subfamily Blastodacninae) caterpillars and a curculionid weevil (Conotrachelus crataegi) was much stronger for R. pomonella larvae infesting the ancestral host hawthorn than the derived host apple. Egg to pupal survivorship was estimated as 52.8% for fly larvae infesting hawthorn fruit without caterpillars and weevils compared to only 27.3% for larvae in harthorns with interspecific insects. Survivorship was essentially the same between fly larvae infesting apples in the presence (44.8%) or absence (42.6%) of interspecific insects. Intraspecific competition among maggots was also stronger in hawthorns than apples. The order or time that a larva exited a hawthorn fruit was a significant determinant of its pupal mass, with earlier emerging larvae being heavier than later emerging larvae. This was not the case for larvae in apples, as the order or time that a larva exited an apple fruit had relatively little influence on its pupal mass. Our findings suggest that decreased performance related to host plant chemistry/nutrition may restrict host range expansion and race formation in R. pomonella to those plants where biotic/ecological factors (i.e. escape from competitors and parasitoids) adequately balance the survivorship equation. This balance permits stable fly populations to persist on novel plants, setting the stage for the evolution of host specialization under certain mitigating conditions (e.g. when mating is host specific and host-associated fitness trade-offs exist).     

Size of orchard trees as a factor affecting behavioural control of apple maggot flies (Dipt., Tephritidae) by traps

Over a 3-year period (1997–1999), we examined the influence of tree size on effectiveness of traps for behaviourally controlling apple maggot flies, Rhagoletis pomonella (Walsh), in small blocks of trees in Massachusetts commercial apple orchards. Traps were red spheres coated with Tangletrap and baited with an attractive synthetic host fruit odour (butyl hexanoate). Traps were placed 6 m apart on perimeter apple trees of each block and were designed to intercept apple maggot flies immigrating into blocks from unmanaged host trees. Based on captures of adults by unbaited red spheres placed near the centre of each block to monitor degree of adult penetration into the interior and on percentages of fruit injured by apple maggot, traps surrounding blocks of small trees (1.5 m canopy diameter) planted at high density were more effective in controlling apple maggot flies (relative to control obtained by insecticide sprays used in comparison blocks) than were traps surrounding blocks of large trees (3.7 m canopy diameter) planted at low density, with traps surrounding blocks of medium-size trees (2.5 m canopy diameter) planted at medium density generally providing an intermediate level of control.     

Trap position and fruit presence affect visual responses of apple maggot flies (Dipt., Tephritidae) to different trap types

Abstract:  We compared responses of apple maggot flies, Rhagoletis pomonella (Walsh), in a commercial apple orchard in Massachusetts with three different types of visual traps baited with synthetic fruit volatiles. The traps were red spheres, Ladd traps (a rectangular yellow panel between two red hemispheres) and rectangular yellow panels. Traps were placed either in optimal position (D rummond , F.; G roden , E.; P rokopy , R. J., 1984: Environ. Entomol. 13, 232–235.) or sub-optimal position (not surrounded by foliage and fruit from all sides). The study was performed over 3 years. Early in the fruiting season, when apples were small and green, red spheres followed by Ladd traps attracted more flies than yellow panels. Ladd traps were the most sensitive to trap positioning, losing effectiveness when placed in sub-optimal position. Late in the fruiting season, in years yielding crops of large red apples, red spheres in optimal position (surrounded by growing red fruit) lost some effectiveness relative to Ladd traps in optimal position. Red spheres in sub-optimal position were not similarly affected, perhaps because of the lack of fruit in the background. Red sphere performance relative to Ladd traps was re-established after removal of competing fruit from trees at harvest. Because Ladd traps are more expensive to purchase and more difficult to position, red spheres are a better option for commercial growers in eastern North America. Conceivably, loss of capturing power of red spheres because of visual competition from red apples towards harvest could be mitigated by adjusting trap positioning.     

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.     

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.     

Uptake of adhesive powders from lure stations by Mediterranean fruit fly (Dipt., Tephritidae)

Powders that are capable of adhering to insect cuticles can act as carrier particles when combined with insecticides, entomopathogens, or pheromones, for targeted insect control. One potential method of delivering the powder to an insect is to lure the insects to stations containing powder using a species‐specific attractant. Here, we report on the uptake of two different powders from lure stations (henceforth called ‘dispensers’) by the Mediterranean fruit fly, Ceratitis capitata, and the transfer of the powders to conspecifics during field studies in Portugal, as part of a research programme to develop lure‐and‐kill technologies based on adhesive powder. Uptake of an electrostatic wax powder, Entostat?, from dispensers was greater than uptake of a proprietary metallic powder, Entomag?, for both wild male C. capitata visiting field‐placed dispensers and laboratory‐reared males confined with dispensers in field cages. In agreement with field data, C. capitata also took up more Entostat than Entomag when artificially dosed on dispenser trays containing powder in the laboratory, and the quantities taken up were shown to be greater than that calculated from field experiments. Increasing the amount of Entostat powder in field‐placed dispensers resulted in greater uptake of powder by visiting male C. capitata. Laboratory‐reared male and female C. capitata were released in field cages in which were hung dispensers containing adhesive powder that were baited with the male attractant trimedlure. After 24 h, the powder was successfully extracted from all males and nearly all females collected, indicating that males probably transferred powder to conspecific females after visiting dispensers. The results underscore that a lure‐and‐kill system based on adhesive powder might have potential for controlling Mediterranean fruit fly and other flying insects.     

Detection of the solanum fruit fly, Bactrocera latifrons (Hendel) in Tanzania (Dipt., Tephritidae)

Abstract:  The presence of the Solanum fruit fly, Bactrocera latifrons , in Africa is reported for the first time, based on trapped and reared specimens in Tanzania. Two new host records, Solanum aethiopicum and Solanum macrocarpon , are reported.     

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.     

Field monitoring of onion maggot (Delia antiqua) fly through improved trapping

Onion maggot (Delia antiqua) is an economically important pest of Allium crops in temperate regions throughout the world. Management of this pest is necessary to achieve economic returns and depends on insecticide regimes and cultural management. Current cultural management especially altering planting date, field location and crop rotation depend on monitoring. We evaluated the effect of shape, size, colour and chemical attractants on trap catch of field populations of adult D. antiqua flies in upstate New York. White, large diameter, spherical traps in conjunction with Delia Lure attractant performed the best in attracting and catching D. antiqua adults. These results suggest an improved means of attracting and capturing D. antiqua populations which could be useful in monitoring efforts and development of attract and kill strategies for pest control.     

Rearing ofChaetorellia australis (Dipt., Tephritidae) on an artificial diet

Larvae of the tephritid flyChaetorellia australis Hering, which infests flowerheads of the yellow starthistleCentaurea solstitialis L. (Asteraceae), and which is a candidate species for biological control, of that weed, were reared for the first time and for two consecutive generations on an artificial diet. At a density of 0.5 eggs per g diet a yield of 18.5% F₁ adults (over hatched eggs), was obtained. When the density of eggs per g diet increased, the yield decreased. At 25°C the duration of development from egg to adult was 20–25 days for both generations. F₁ adults reared on artificial diet had an average longevity of 32.0 days for males and 34.9 days for females. The fecundity was 53.3 eggs per female and the egg hatchability 88.8%. These and other biological parameters are similar to those of wild flies.      

Commercial Yellow Sticky Strips more attractive than yellow boards to western cherry fruit fly (Dipt., Tephritidae)

Bright yellow sticky rectangles made of paper boards were previously identified as the most effective traps for capturing western cherry fruit fly, Rhagoletis indifferens Curran (Dipt., Tephritidae). However, no data on the effectiveness of commercial sticky yellow plastic traps against R. indifferens have been reported. In tests conducted in sweet cherry trees [Prunus avium (L.) L.] in Washington state (USA) using ammonium carbonate as the chemical lure, commercial plastic ‘Yellow Sticky Strips’ made of translucent high‐impact polystyrene captured ~two or three times more flies than commercial sticky yellow‐folded Pherocon^® AM and Alpha Scents boards. Yellow Sticky Strips also minimized captures of non‐target flies and bees per surface area compared with Pherocon^®AM and/or Alpha Scents boards. Trap size and adhesive type were not factors for greater catches of R. indifferens. However, more flies were caught on the shade‐facing side of Yellow Sticky Strips, which was brightly illuminated, than on the shade‐facing side of boards, which was darker, suggesting differential light passage was a factor. The Yellow Sticky Strips could be very useful for monitoring R. indifferens in detection programmes and based on the results of this study can replace yellow boards. They are also useful because they are relatively unattractive to non‐target insects.     

Mediterranean fruit fly (Dipt., Tephritidae) suppression in persimmon through bait sprays in adjacent coffee plantings*

Abstract: Oriental persimmon, Diospyros kaki L., in Upper Kula on the island of Maui (Hawaii) is attacked by the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). Recent suppression trials using mass trapping with a synthetic food‐based bait, initiated in alternate host crops before the start of persimmon season, had shown promise as a means of reducing C. capitata population levels. However, this did not adequately suppress C. capitata population where there were adjacent plantings of coffee, Coffea arabica L., a favoured alternate host, which bears fruits before and during the persimmon season. To improve C. capitata population suppression, we applied a spinosad‐based bait spray to coffee plants, starting before persimmon fruits became susceptible to oviposition by the Mediterranean fruit fly. The bait spray suppressed the C. capitata population and led to reduced infestation of both coffee cherries and persimmon fruits. Percentage parasitization of C. capitata in coffee cherries by established biological control agents, primarily Fopius arisanus (Sonan), was not significantly different in unsprayed vs. sprayed plots even after 11 weekly sprays. These results suggest that mass trapping, combined with spinosad‐based bait sprays, are control components that are compatible with biological control and can be combined in an integrated pest management system for C. capitata.     

Seasonality and host utilization of the invasive fruit fly, Bactrocera invadens (Dipt., Tephritidae) in central Tanzania

Abstract:  The temporal occurrence of the invasive and economically important pest fruit fly, Bactrocera invadens was studied in three agro-ecological areas of Morogoro Region, central Tanzania, during 2004–2005. Weekly and monthly trappings were carried out with methyl eugenol, protein bait and synthetic food attractant. Bactrocera invadens was permanently present at low and mid-altitudes (380–520 m a.s.l.) with peak periods coinciding with the fruiting season of mango ( Mangifera indica ) and guava ( Psidium guajava ). At high altitude (1650 m a.s.l.) its incidence was only temporal and apparently the result of dispersal from lower altitudes after the mango fruiting season. Rearing results showed mango, loquat ( Eriobotrya japonica ), guava and grapefruit ( Citrus  ×  paradisi ) to be the favoured commercial host fruits. Other Citrus species, cucurbits, papaya ( Carica papaya ) and avocado ( Persea americana ) were less favoured.     

Egg–larval opiine parasitoids (Hym., Braconidae) of tephritid fruit fly pests do not attack the flowerhead-feeder Trupanea dubautiae (Dipt., Tephritidae)

Abstract:  We investigated the potential impact of three opiine tephritid fruit fly parasitoids: Fopius arisanus (Sonan), Fopius caudatus (Szépligeti) and Fopius ceratitivorus Wharton, on the non-target native Hawaiian tephritid, Trupanea dubautiae (Bryan), infesting flowerheads of the endemic Asteraceae shrub Dubautia raillardioides Hillebrand. The three species are the only known opiine fruit fly parasitoids that attack host eggs (but occasionally attack first instars). F. arisanus , which originated in southeast Asia, is now widely established in Hawaii and elsewhere in the world, while the other two are African species currently in quarantine in Hawaii. In the laboratory, field-collected flowerheads of D. raillardioides containing T. dubautiae eggs and first instar larvae were exposed to naïve female wasps of each of the three Fopius species in the absence (no-choice test) or presence (choice test) of papaya fruit infested with eggs of the Mediterranean fruit fly Ceratitis capitata (Wiedemann), the parasitoids' normal host. All three Fopius species visited the papaya fruit much more often than the flowerheads, and displayed normal ovipositional responses (probing and stinging) on the fruit in the choice test. None of the three parasitoid species showed ovipositional responses to flowerheads in either choice or no-choice tests. As a result, not a single T. dubautiae egg or larva was attacked by any of the three parasitoids, indicating that these parasitoids of pest tephritids are unable to recognize the microhabitats of flowerhead-feeding tephritids. The results suggest minimal risk of non-target impact in these biological control agents.     

Fruit trap: A detection and collection tool for opiine parasitoids (Hym.: Braconidae) of the oriental fruit fly,Bactrocera dorsalis (Dipt.: Tephritidae)

A fruit trap was developed for detection and collection of the opiine parasitoids of the oriental fruit fly,Bactrocera (=Dacus)dorsalis (Hendel). Gravid females ofBiosteres arisanus (Sonan), an egg-larval parasitoid, orDiachasmimorpha longicaudata (Ashmead) andPsytallia incisi (Silvestri), both larval parasitoids, were lured to parasitize the eggs or larvae ofB. dorsalis inoculated in ripe papaya fruits,Carica papaya L. Progenies ofB. arisanus were consistently recovered from papaya fruits inoculated withB. dorsalis eggs (subsequently referred to as egg fruit traps). Except in Moloaa on Kauai (6%), higher percentage ofB. dorsalis parasitization (range=38–43%) was recorded in Hilo, island of Hawaii and Waimanalo and Poamoho, island of Oahu. Progenies ofD. longicaudata and a fewP. incisi were recovered from papaya fruits artificially infested withB. dorsalis larvae (subsequently referred to as larval fruit traps). The recovery of parasitoid progenies from larval fruit traps suspended from papaya trees did not differ significantly from larval fruit traps placed on the ground. In both methods of trap placement, percent parasitization ofB. dorsalis byD. longicaudata (predominant species) ranged from 58–60%. On the other hand, significantly moreB. arisanus thanD. longicaudata andP. incisi adults (larval parasitoids) were recovered from fully ripened to highly deteriorated papaya fruits collected from papaya trees or ground (fallen fruits).     

Evaluation of border sprays for managing the codling moth (Tortricidae: Lepidoptera) and the apple maggot (Tephritidae: Diptera) in Ontario apple orchards

The efficacy of two insecticide control programs for managing the codling moth, Cydia pomonella (L.), and the apple maggot, Rhagoletis pomonella (Walsh), were compared in the Georgian Bay, London, Niagara, and Quinte apple production areas of Ontario during 1995, 1996, and 1997. In the border spray program, an initial cover spray of organophosphorus insecticide was applied to eradicate codling moths that may have colonized a test plot during the previous growing season. Subsequent sprays were applied only to a four-tree-wide zone (approximately 20 wide) around the perimeter of the plot to control immigrating codling moths or apple maggots. In the cover spray program, all sprays of organophosphorus insecticide were applied to the entire plot. Apple maggot injury was significantly greater in border spray program plots than in cover spray program plots only during 1995 in the London production area. There was no significant difference in codling moth injury between border spray and cover spray plots in the four production areas during the three-year study. The elimination of cover sprays from border spray plots during July and August may have left the apple crop more susceptible to damage by second generation larvae of the obliquebanded leafroller, Choristoneura rosaceana (Harris), in the London production area during 1995. There was a trend of increasing codling moth injury from 1995 to 1997 in two border spray plots, and apple maggot injury was detected in these plots during the third year of the study.     

The efficacy of an improved form of the mass-trapping method, forthe control of the olive fruit fly, Bactrocera oleae (Gmelin) (Dipt., Tephritidae): pilot-scale feasibility studies

Abstract:  The efficacy of an improved form of the mass-trapping method for the control of the olive fruit fly, Bactrocera oleae (Gmelin) was tested for 4 years in a pilot test at Tanagra Voeotia, Greece. Improvements consisted of the extension of the active life of the toxic trap used, active life referring both to its attracting and killing properties, as well as in trap deployment, which combined efficacy and low cost. The method was compared to bait sprays applied from the ground, which constitutes the current standard method for the control of this pest. Both pest population density and fruit infestation levels, the main parameters used for the evaluation of the two methods were considerably lower during all 4 years of tests in the orchards protected by mass trapping compared with those in the orchards protected by bait sprays. Furthermore no complementary measures were required in the mass-trapping orchards for acceptable crop protection, which was not the case under certain conditions, prior to the introduction of the recent improvements. The cost of the mass-trapping method was approximately US$ 0.40 per tree per year compared with US$ 0.35 for bait sprays (figures of the Greek Ministry of Agriculture). However, the mass-trapping method reduces the amount of insecticide used for olive protection by 99.5% (15 mg a.i. per tree per year as opposed to 3 g in the case of bait sprays). A considerable reduction in the cost of the mass-trapping method is expected with the extension of its use and the mass production of materials used, especially traps.     

Evaluation of synthetic food-based attractants for female Mediterranean fruit flies (Dipt., Tephritidae) in McPhail type traps

During summer 1997 field experiments were conducted on the island of Chios, Greece, to compare captures of female Mediterranean fruit flies (medflies), Ceratitis capitata (Wiedemann), in traps baited with either synthetic female-targeted lures or a standard protein bait (NuLure and borax). The synthetic lures contained ammonium acetate,1,4 diaminobutane (putrescine), and trimethylamine. Two trap types (International Pheromone's McPhail Trap (IPMT) and the Tephri trap) were tested as either wet or dry. Wet IPMT traps baited with the synthetic attractants were the most attractive of all trap combinations tested and captured 2.1 times more female medflies and 1.8 times more total medflies than traps baited with NuLure and borax. Traps containing the synthetic attractant captured approximately 4.6 times fewer nontarget insects than NuLure baited traps. Vapona used in IPMT traps was repellent to medflies and dry traps with lower concentrations of Vapona were approximately 1.5 times less attractive to female medflies than traps containing water. Even with a decrease in attractiveness, the dry traps were significantly more effective for females and more practical for mass trapping and monitoring than the currently used traps baited with protein solutions.