Integration of insecticidal, phagostimulatory, and visual elements of an attract and kill system for apple maggot fly (Diptera: Tephritidae).

The apple maggot fly, Rhagoletis pomonella (Walsh), is a key pest of apple in eastern North America that has been historically controlled with organophosphate insecticide applications. Here we report on progress toward development of an effective and maintenance-free attracticidal sphere system for this pest species. In our studies, we evaluated lethality of spinosad in combination with a feeding stimulant (sucrose) to replace a Tangletrap sticky coating as the killing agent of a sphere-based behavioral control system. Spinosad was incorporated into cylindrical and contoured controlled-release caps that were fixed atop visually stimulating sphere bases. For both cap styles, spinosad at or near 1.0% (a.i.) proved to be a relatively durable fly-killing agent, providing approximately equal to 85% mortality after simulated rainfall exposure reflective of average season-long precipitation levels experienced during the primary period of risk of apple maggot injury to fruit in the northeastern United States. In field trials, we assessed the impact of color degradation of contoured controlled-release caps on visual responsiveness of apple maggot fly and found that it had no significant impact on captures. In commercial orchard trials designed to evaluate the potential of attracticidal spheres with contoured caps for direct control of apple maggot, a perimeter-based deployment provided protection comparable to plots receiving 1-2 whole-plot insecticide applications.

Effectiveness of odor-baited trap trees for plum curculio (Coleoptera: Curculionidae) monitoring in commercial apple orchards in the northeast.

The plum curculio, Conotrachelus nenuphar (Herbst), is a key pest of pome and stone fruit in eastern and central North America. For effective management of this insect pest in commercial apple (Malus spp.) orchards in the northeastern United States and Canada, one of the greatest challenges has been to determine the need for and timing of insecticide applications that will protect apple fruit from injury by adults. In a 2004-2005 study, we assessed the efficacy and economic viability of a reduced-risk integrated pest management strategy involving an odor-baited trap tree approach to determine need for and timing of insecticide use against plum curculio based on appearance of fresh egg-laying scars. Evaluations took place in commercial apple orchards in seven northeastern U.S. states. More specifically, we compared the trap-tree approach with three calendar-driven whole-block sprays and with heat-unit accumulation models that predict how long insecticide should be applied to orchard trees to prevent injury by plum curculio late in the season. Trap tree plots received a whole-plot insecticide spray by the time of petal fall, and succeeding sprays (if needed) were applied to peripheral-row trees only, depending on a threshold of one fresh plum curculio egg-laying scar out of 25 fruit sampled from a single trap tree. In both years, level of plum curculio injury to fruit sampled from perimeter-row, the most interior-row trees and whole-plot injury in trap tree plots did not differ significantly from that recorded in plots subject to conventional management or in plots managed using the heat-unit accumulation approach. The amount of insecticide used in trap tree plots was reduced at least by 43% compared with plots managed with the conventional approach. Advantages and potential pitfalls of the bio-based trap tree approach to plum curculio monitoring in apple orchards are discussed.

Odor-baited trap trees: a novel management tool for plum curculio (Coleoptera: Curculionidae).

The plum curculio, Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae), one of the most important pests of apple (Malus spp.) in eastern and central North America, historically has been managed in New England apple orchards by three full block insecticide applications. Efforts to reduce insecticide inputs against plum curculio include perimeter row sprays, particularly after petal fall, to control immigrating adults. The odor-baited trap tree approach represents a new reduced input strategy for managing plum curculio based on the application of insecticides to a few perimeter-row trap trees rather than the entire perimeter row or full orchard block. Here, we compared the efficacy of a trap tree approach with perimeter row treatments to manage populations after petal fall in commercial apple orchards in 2005 and 2006. Injury was significantly greater in trap trees compared with unbaited perimeter row treated trees in both years of the study. In 2005, heavy rains prevented growers from applying insecticide applications at regular intervals resulting in high injury in nearly all blocks regardless of type of management strategy. In 2006, both the trap-tree and perimeter-row treatments prevented penetration by immigrating populations and resulted in economically acceptable levels of injury. The trap tree management strategy resulted in a reduction of approximately 70% total trees being treated with insecticide compared with perimeter row sprays and 93% compared with standard full block sprays.

3 using odor-baited trap trees as sentinels to monitor plum curculio (Coleoptera: Curculionidae) in apple orchards.

In commercial orchards in Massachusetts in 2003, we conducted experiments aimed at developing guidelines for use of perimeter-row trap trees baited with grandisoic acid plus benzaldehyde as sentinels in a practical approach to determining need and timing of insecticide applications against overwintered plum curculios, Conotrachelus nenuphar (Herbst). Evaluations were based on percentages of sampled fruit injured by plum curculio. Trap trees baited with grandisoic acid released at approximately 1 mg/d plus benzaldehyde released at approximately 40 mg/d performed as well as or better than trap trees baited with greater or lesser amounts of these attractants in combination. The distance over which a trap tree baited with such odor was effective in aggregating damage to fruit extended to at least 31-33 m (maximum evaluated) along a perimeter row. Trap trees at corners of orchard blocks were as effective as perimeter-row trap trees midway between corner trees. Within the canopy of a trap tree, damage did not tend significantly to be localized in the vicinity of the odor source but tended to be rather evenly distributed among various sectors of the canopy. Finally, among three candidate thresholds evaluated as a trigger for insecticide application, a threshold of one freshly injured fruit proved better than thresholds of two or four freshly injured fruit out of 50 fruit sampled on a trap tree in assuring that orchard-wide damage would remain below a preset economic injury level of 1%. Our findings lead us to suggest that after a whole-orchard application of insecticide to apple trees in Massachusetts orchards shortly after petal fall, subsequent applications of insecticide against plum curculio can be confined to peripheral-row trees and be driven by a provisional threshold of one freshly injured fruit out of 50 fruit sampled on a perimeter-row trap tree baited with the above-mentioned odor.

Effectiveness of GF-120 fruit fly bait spray applied to border area plants for control of melon flies (Diptera: Tephritidae).

In a field study in Hawaii, color-marked protein-deprived and protein-fed female melon flies, Bactrocera cucurbitae Coquillett, were released within canopies of unsprayed sorghum plants (a nonhost of melon flies) outside of a border area of unsprayed or bait-sprayed sorghum plants or open space that surrounded cucumbers, a favored host of melon flies. Application of bait spray to sorghum or sugarcane surrounding host plants of melon flies is a common practice for melon fly control in Hawaii. GF-120 Fruit Fly Bait spray proved very effective in preventing protein-deprived females from alighting on cucumbers (23% of released females were observed dead on bait-sprayed sorghum; 0% were observed alive on cucumbers), but proved less effective in suppressing protein-fed females (14% of released females were observed dead on bait-sprayed sorghum; 11% were observed alive on cucumbers). No females were found dead on unsprayed sorghum. Compared with open space surrounding cucumbers, the presence of unsprayed sorghum as surrounding border area neither significantly enhanced nor significantly inhibited the ability of either type of female with respect to finding cucumbers. Greenhouse cage assays revealed that compared with droplets of water, droplets of GF-120 Fruit Fly Bait spray were highly attractive to protein-deprived females within 1 h of bait spray application to sorghum, but lost about half of their attractiveness within 5 h and all of it within 24 h under the dry greenhouse conditions used for maintaining baited-sprayed sorghum plants in these assays. Laboratory cup assays showed that bait spray droplets remained highly toxic to protein-deprived females 24 h after application, but lost nearly half of their toxicity within 4 d under laboratory exposure and nearly all of it after approximately 8 mm of rainfall. Combined findings suggest that application of GF-120 Fruit Fly Bait spray to nonhost plants for melon fly control either be made often enough to overcome loss of attractiveness of bait spray droplets to females or that bait spray be applied to nonhost plants that are themselves attractive to the females.

Odor-baited trap trees: a new approach to monitoring plum curculio (Coleoptera: Curculionidae).

We compared a trap approach with a trap-tree approach to determine the need and timing of insecticide applications against overwintered adult plum curculios, Conotrachelus nenuphar (Herbst.), in commercial apple orchards in Massachusetts in 2002. All traps and trap trees were baited with benzaldehyde (attractive fruit odor) plus grandisoic acid (attractive pheromone). Sticky clear Plexiglas panel traps placed at orchard borders, designed to intercept adults immigrating from border areas by flight, captured significantly more adults than similarly placed black pyramid traps, which are designed to capture adults immigrating primarily by crawling, or Circle traps wrapped around trunks of perimeter-row trees, which are designed to intercept adults crawling up tree trunks. None of these trap types, however, exhibited amounts of captures that correlated significantly with either weekly or season-long amounts of fresh ovipositional injury to fruit by adults. Hence, none appears to offer high promise as a tool for effectively monitoring the seasonal course of plum curculio injury to apples in commercial orchards in Massachusetts. In contrast, baiting branches of selected perimeter-row trees with benzaldehyde plus grandisoic acid led to significant aggregation (14-15-fold) of ovipositional injury, markedly facilitating monitoring of the seasonal course of injury to apples. A concurrent experiment revealed that addition of other synthetic fruit odor attractants to apple trees baited with benzaldehyde plus grandisoic acid did not enhance aggregation of ovipositional injury above that of this dual combination. We conclude that monitoring apples on odor-baited trap trees for fresh ovipositional injury could be a useful new approach for determining need and timing of insecticide application against plum curculio in commercial orchards.

Field evaluation of plant odor and pheromonal combinations for attracting plum curculios.

The attractiveness of different synthetic host odors and a synthetic aggregation pheromone (grandisoic acid [GA]) to overwintered adult plum curculios (PCs), Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae), was examined using two types of traps (sticky panels and black pyramids) placed in border areas surrounding an unsprayed section of an apple orchard in Massachusetts. In 2001, we evaluated the response of PCs to three synthetic fruit volatiles (benzaldehyde [BEN], ethyl isovalerate [EIV], and limonene [LIM]) assessed alone and in combination with GA, as well as the response to GA alone and a no-odor (control) treatment. BEN was the only host volatile that synergized the response of PCs to GA for both trap types. For both trap types, GA was as attractive to PCs as a single component as when in combination with either EIV or LIM. In 2002, four release rates of BEN (0, 2.5, 10, and 40 mg/day) and two release rates of GA (1 and 2 mg/day) were evaluated for attractiveness to PCs using panel and pyramid traps. For panel traps, an increase in amount of GA released (from 1 to 2 mg/day) was associated with a 35% increase in captures. However, PC captures by pyramid traps were similar regardless of the amount of GA released. For panel traps, 10 and 40 mg/day of BEN were the most attractive release rates regardless of the amount of GA released. For pyramid traps baited with GA. PC captures were enhanced by the presence of BEN, regardless of release rate. In 2003, GA at 1 mg/day + BEN at 80 mg/day of release did not enhance PC captures by panel traps relative to lower release rates of BEN. Pyramid traps releasing GA at 1 mg/day performed best when baited with BEN at 10 mg/day of release; a release rate of 80 mg/day of BEN decreased the attractiveness of the binary combination of BEN + GA. Combined results suggest that BEN at 10 mg/day + GA at 1 mg/day of release constitutes an attractive lure that may improve the effectiveness of monitoring traps for PCs.

Effect of physiological and experiential state ofBactrocera tryoni flies on intra-tree foraging behavior for food (bacteria) and host fruit.

Using caged host trees on which we manipulated food and oviposition sites, we investigated the foraging behavior of individually-releasedBactrocera tryoni (Diptera: Tephritidae) females in relation to state of fly hunger for protein, presence or absence of bacteria as a source of protein, degree of prior experience with host fruit, and quality of host fruit for oviposition. One aim was to evaluate whether it is immature or matureB. tryoni females that are responsible for initially inoculating host fruit surfaces with "fruit-fly-type" bacteria, the odor of which is known to attractB. tryoni females. We found that 3-week-old immature females provided with sucrose but deprived of protein from eclosion had a much greater propensity than 3-week-old protein-fed mature females to visit vials containing fruit-fly-type bacteria, irrespective of whether vials were associated with adjacent host fruit or not. In the absence of associated bacteria in vials, immature females had a much lower propensity than mature females to visit host fruit. In the presence of bacteria in vials, however, propensity of immature and mature females to visit fruit was about equal. Mature (but not immature) females were more inclined to visit fruit that ranked higher for oviposition (nectarines) than fruit that ranked lower (sweet oranges). Mature females that attempted oviposition during a single 3-min exposure period to a nectarine prior to release were much more likely to find a nectarine than were mature females naive to fruit or immature females with or without prior contact with fruit. Exposure to a nectarine before release did not affect the propensity of either mature or immature females to alight on an odorless visual model of a nectarine, however. As judged by numbers of leaves visited, protein-deprived immature females were more active than protein-fed mature females, irrespective of the sorts of resources on a tree. Together, our findings lead us to conclude that (1) the firstB. tryoni females to arrive on the fruit of a host tree and therefore inoculate the fruit with fruit-fly-type bacteria are unlikely to be sexually immature, but to be mature as a result of having earlier acquired protein elsewhere, (2) the odor of colonies of fruit-fly-type bacteria when associated with host fruit will attract protein-hungry but not protein-fed females, and (3) the odor of the fruit itself will attract mature females (especially experienced ones) but not immature females. These findings illustrate the value of considering jointly the state of a resource patch together with the physiological and experiential state of the individual when investigating the foraging behavior of an insect.

Behavioral evidence for host races in Rhagoletis pomonella flies.

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.

The effect of prior adult experience on components of habitat preference in the apple maggot fly (Rhagoletis pomonella).

Numerous authors have suggested that genetic subdivision within a population in a heterogeneous environment is more likely if individuals tend, through prior experience, to breed in the same habitat in which they developed. Under semi-field conditions we demonstrate that prior adult experience alters habitat preference in the apple maggot fly, Rhagoletis pomonella (Tephritidae), a frugivorous parasitic insect thought to have undergone sympatric divergence in host use in historical times. Females exposed to a particular host fruit species - apple (Malus pumila) or hawthorn (Crataegus mollis) - in a field cage oviposited at a higher rate in test fruit of that species than did inexperienced females or females exposed to the other species. Females exposed to a particular host fruit species also tended to remain longer in test trees harboring fruit of that species than did inexperienced females or females exposed to the other species. Prior adult experience thus alters two components of habitat preference in the apple maggot fly: oviposition preference and habitat fidelity. We discuss how these effects of experience on habitat preference should increase the likelihood that individuals mate assortatively and may further increase the likelihood that apple maggot populations become genetically subdivided.

Host visitation sequence as a determinant of search persistence in fruit parasitic tephritid flies.

We tested, through field experiments and simulation models, the hypothesis that fruit-searching tephritid fruit flies adjust their within-tree search persistence according to the sequence and timing of encounters with parasitized (i.e. egg-infested) and unparasitized Crataegus sp. host fruit. In the field, we presented flies with 4 different sequences of unparasitized [=C] and parasitized [=M] fruit: 5M+0C; 1C+5M; 5M+1C, 1C+0M. Following fruit presentation, flies were permitted to forage freely within trees, which harboured no fruit, until emigration occurred. Under these conditions, flies that encountered the aforementioned different sequences of hosts, displayed differences in Giving Up Time, measured as active foraging time and number of leaf visits, in a manner predicted prior to testing. These differences were, however, not statistically significant. Based upon the results described above, we then built 3 simulation models that predicted within-tree Giving Up Times for individual flies: Model 1-Giving Up Time is incremented and decremented by fixed amounts following encounters by the fly with suitable (i.e. for oviposition) and unsuitable hosts, respectively; Model 2-similar to Model 1, but increment and decrement values are variable and are dependent upon the time since previous encounters by the fly with suitable and unsuitable hosts; Model 3-Giving Up Time is fixed. Comparison with previously reported field data for tephritid flies showed that Model 2 predicted rather well, and significantly better than Models 1 and 3, Giving Up Time for wild type tephritid flies under seminatural field conditions. We discuss our results in light of contemporary foraging theory.