Leaf color used by cabbage root flies to distinguish among host plants.

In experiments in which spectrophotometric reflectance patterns of real leaves were mimicked with mixtures of artists' pigments, leaf color was shown to be a character used by cabbage root flies, before landing on leaves, to discriminate among the host plant cultivars radish, green cabbage, and red cabbage. It may be possible to take advantage of factors that affect leaf color, such as epicuticular bloom, pubescence, and masking of chlorophyll by other pigments, to decrease the attraction of certain pest insects to plants.

Associative learning in egglaying site selection by apple maggot flies.

Evidence is presented demonstrating that associative learning during oviposition in Crataegus or apple hosts can significantly influence the propensity of apple maggot flies to accept or reject these hosts in future encounters. The data suggest that within resource patches of a given host type there may be an enhancement of foraging efficiency.

Controlled release of sugar and toxicant from a novel device for controlling pest insects.

A novel biodegradable device, designed for long-lasting residual effectiveness of feeding stimulant (sugar) and insecticide (dimethoate) against apple maggot files and other insects, was formulated. The device is an 8-cm diameter fruit-mimicking sphere, consisting of 42-50% sugar entrapped in a mixture of gelatinized corn flour and wheat flour in the presence of glycerin, and coated with a layer of latex paint containing dimethoate and sugar. We found that the outer layer of paint prevents cracking of the sphere upon drying and creates a barrier to control the release of both sugar and dimethoate. Releases of each ingredient were screened first by chemical analysis and then by bioassays in the laboratory and in field cages against apple maggot flies. Chemical analysis demonstrated strong potential for controlled release of water-soluble feeding stimulant and water-insoluble insecticide measured as a function of the amount of rainfall and duration of exposure time. Field results showed greater than 70% insecticidal activity after 11 weeks of sphere exposure in an orchard. This device has the potential to be used for a variety of insect-control applications through manipulating its shape, color and texture into forms known to be attractive to target insects, and by employing various toxicants designed to be effective against such insects.

Sex differences in movement between natural feeding and mating sites and tradeoffs between food consumption, mating success and predator evasion in Mediterranean fruit flies (Diptera: Tephritidae).

Systematic quantitative observations of the location and diel pattern of adult Mediterranean fruit fly, Ceratitis capitata (Wiedemann), activities were carried out in an orange grove and surroundings on the island of Chios in Greece. Natural fly foods were assessed for their contribution to fly longevity, fecundity and fertility. There were diel shifts in male and female location. Females required a substantial and varied diet to realize peak fecundity. This diet was acquired away from the primary host, orange. Foraging for food throughout most of the day on fig and non-host foliage (including feeding on bird droppings) as well as on fig fruit and grapes, females dispersed and fed more than males. A diet of grapes alone did not support any fecundity, contributing only to longevity. A diet of figs alone, on the other hand, sustained both longevity and egg production. Bird feces alone supported neither egg production nor longevity. However, when added to a diet of figs, bird feces significantly increased fly fecundity. Throughout most of the day, males aggregated in leks within the inner canopy of the primary host, orange. The arrival here during the warmest hours of the day of receptive females, followed by pair formation, reinforced the lek mating system on host foliage. In the afternoon, females shifted to orange fruit where they suffered from high predation mortality while ovipositing. Soon after, males also shifted to orange fruit, where they attempted matings with non-receptive ovipositing females. Male feeding on fig fruit occurred late in the day, a time when they were least likely to find a mate. Male survival did not differ between the natural diets. Tradeoffs between food consumption, mating success and predator evasion are discussed for each sex and related to fruit fly mating systems.

Response of plum curculio (Coleoptera: Curculionidae) to odor-baited traps near woods.

Response of overwintered plum curculios, Conotrachelus nenuphar (Herbst), to odor-baited traps was evaluated from the beginning until nearly the end of emigration from overwintering sites in woods. We evaluated clear sticky Plexiglas panels and black pyramid traps placed close to woods adjacent to apple trees in an unsprayed section of an orchard. Traps were baited with aggregation pheromone (grandisoic acid) alone or in combination with one of six synthetic fruit volatiles (benzaldehvde, decyl aldehyde, E-2-hexenal, ethyl isovalerate, hexyl acetate, or limonene). Unbaited traps served as a control treatment. Plum curculio emigration from woods was divided into early-, mid-, and late-season periods based primarily on phenological stage of apple bud and fruit development (tight cluster to bloom, petal fall, and fruit set, respectively). During both early- and late season, panel and pyramid traps baited with benzaldehyde plus pheromone were significantly more attractive than any other traps (baited or unbaited), except panel traps baited with ethyl isovalerate plus pheromone in early season, which likewise captured significantly more adults than unbaited panel traps. During midseason, no lures were significantly attractive, possibly due to prevailing cool weather, unfavorable for adult activity. Over the entire season, panel or pyramid traps baited with benzaldehyde plus pheromone captured nearly six times as many plum curculios as unbaited traps of each type, whereas traps baited with pheromone alone captured about twice as many as unbaited traps of each type. We provide information on sex ratio, female maturity stage, and mating status, and several weather parameters associated with trap captures. We conclude that panel or pyramid traps, or a combination, baited with benzaldehyde plus pheromone placed at borders of plum curculio overwintering sites can be a valuable tool for monitoring the beginning, peak, and end of adult immigration into apple orchards.

Toxicity and residual effectiveness of insecticides on insecticide-treated spheres for controlling females of Rhagoletis pomonella (Diptera: Tephritidae).

This study evaluated the toxicity of five technical-grade insecticides of four different classes to apple maggot females, Rhagoletis pomonella (Walsh), following a 10-min exposure period in insecticide-coated glass jars, with or without a feeding stimulant (sucrose) present. According to LC90 values for toxicity by ingestion and tarsal contact, imidacloprid was 1.5 times more toxic than dimethoate or abamectin, diazinon was less toxic, and phloxine B (a phototoxic dye) least toxic. Based on LC90 values for tarsal contact alone, dimethoate was 2.3, 4.0, and 18.4 times more toxic than imidacloprid, abamectin, and diazinon, respectively. Contact alone with phloxine B caused no mortality. When exposure was assessed using spheres coated with a latex paint mixture containing sucrose and formulated dimethoate (Digon 400 EC) or imidacloprid (Provado 1.6 F) at concentrations ranging from 5 to 70 g (AI)/cm2, both insecticides showed reduced effectiveness compared with toxicities from glass jar tests, with Digon two times more toxic than Provado. After exposure to artificial rainfall and retreatment with sucrose, Digon- and Provado-treated spheres exhibited greatest residual effectiveness, with diazinon-treated spheres less effective. Spheres treated with formulated abamectin (Agri-Mek 0.15 EC) at 1.0% (AI) performed only slightly better than phloxine B-treated spheres, which completely lost effectiveness after exposure to rainfall. Spheres treated with formulated imidacloprid (Merit 75 WP) at 1.5% (AI) showed equal or better residual efficacy in killing apple maggot flies (> 80% mortality, shorter lethal duration of feeding) over a 12-wk exposure period to outdoor weather than spheres treated with Digon at 1.0% (AI) after both types were retreated with sucrose. Our results indicate that imidacloprid is a promising safe substitute for dimethoate as a fly killing agent on lure-kill spheres. Imidacloprid formulated as Merit 75 WP had greater residual efficacy than imidacloprid formulated as Provado 1.6 F.

Comparison of neonicotinoid insecticides for use with biodegradable and wooden spheres for control of key Rhagoletis species (Diptera: Tephritidae).

Field-based studies and laboratory bioassays were conducted with apple maggot, Rhagoletis pomonella (Walsh), and blueberry maggot, Rhgoletis mendax Curran, flies to investigate the performance and duration of activity of insecticide-treated biodegradable and wooden spheres for control of Rhagoletis species. Four neonicotinoid insecticide treatments including imidacloprid, thiamethoxam, and thiocloprid at 2% (AI) were evaluated with biodegradable spheres. In 1999, significantly more apple maggot flies were found killed by imidacloprid-treated spheres compared with thiamethoxam-treated spheres during early and late season. In 2000, spheres treated with either of two formulations of imidacloprid killed significantly more apple maggot flies compared with thiamethoxam, thiocloprid, and untreated spheres. In blueberries, there were no significant differences between the numbers of blueberry maggot flies killed by both imidacloprid-treated or thiamethoxam-treated spheres in 1999. However, during the 2000 blueberry field season, both formulations of imidacloprid were significantly more effective in killing blueberry maggot flies compared with spheres treated with thiamethoxam, thiocloprid and untreated controls. Overall, spheres treated with thiocloprid were ineffective and did not kill significantly more apple maggot or blueberry maggot flies compared with the controls. Laboratory bioassays showed that the effectiveness of field-exposed spheres treated with imidacloprid at 4 and d 8% (AI) and thiamethoxam at 4% (AI) in killing apple maggot flies was not significantly reduced over a 12-wk aging period. Additionally, wooden spheres aged outdoors for 12 wk with and without mold maintained residual activity in laboratory tests, whereas biodegradable spheres of equal aging, with and without mold lost their effectiveness in killing apple maggot flies. In other studies, we confirmed that the addition of an external feeding stimulant (sucrose) significantly increases the effectiveness of both biodegradable and wooden spheres treated with imidacloprid at 2% (AI).

Attraction of adult Rhagoletis indifferens (Diptera: Tephritidae) to unbaited and odor-baited red spheres and yellow rectangles.

Five sizes of red spheres (4, 6, 8, 10, and 12 cm diameter) and 2 orientations of yellow rectangles (vertical and V) were evaluated as unbaited sticky-coated traps for western cherry fruit flies, Rhagoletis indifferens Curran, in unmanaged cherry trees in Washington and Oregon. Red spheres that were 10 cm in diameter attracted more flies than red spheres that were 8 or 12 cm in diameter and significantly more flies than 4- or 6-cm spheres and yellow rectangles of either orientation. In a 2nd test, red spheres (10 cm diameter) baited with ammonium carbonate alone or ammonium carbonate plus putrescine attracted significantly more R. indifferens than similar spheres baited with ammonium acetate alone, putrescine alone, 3-methyl-1-butanol alone, or combinations of these substances. In a 3rd test, vertical yellow rectangles baited with ammonium carbonate alone attracted numerically more R. indifferens than any of the aforementioned substances alone or in combination. We discuss the potential value of 10-cm red spheres baited with ammonium carbonate for monitoring and direct control of R. indifferens.

Attraction ofCeratitis capitata (Diptera: Tephritidae) flies to odor of coffee fruit.

On potted nonfruiting host trees in outdoor field cages, we evaluated attraction of released mature laboratory-cultured or wild-origin Mediterranean fruit flies (medflies) to odor of freshly picked fruit of host and nonhost plants. Odor of ripe intact or crushed coffee fruit (the presumed ancestral host of medflies) was significantly more attractive than odor of ripe intact or crushed fruit of five lower-ranking hosts and three nonhosts. Odor of crushed coffee fruit was significantly more attractive than odor of intact coffee fruit. Odor of ripe or near-ripe coffee fruit was significantly more attractive than odor of unripe coffee fruit. Immature females (without eggs) were significantly more attracted to odor of a proteinaceous food lure than to odor of ripe coffee fruit, whereas the reverse was true for mature females carrying a high egg load. In some trials, males proved as discriminating as females in favor of coffee fruit odor, but in several other trials males were less discriminating than females. Response patterns of mature laboratory-cultured females were similar to those of mature wild-origin females. In a field of coffee plants, attraction of natural-population females was significantly greater to odor of ripe coffee fruit than to water but was not greater than attraction to odor of proteinaceous food. Findings are discussed in relation to potential use of synthetic volatiles of coffee or other host fruit in traps for monitoring or controlling medflies.

Joining and avoidance behavior in nonsocial insects.

Groups of two or more consexual conspecific adults of many kinds of nonsocial insects have been observed to form at feeding, mating, ovipositional, or sheltering sites. Conversely, adults of these same insects have been observed to avoid joining consexual conspecifics (or their progeny) and to place themselves (or their progeny) at some distance that results in spacing. Examples from various taxa illustrate that mechanisms underlying joining or avoidance behavior differ among species, as do types of benefits and costs to individuals who decide to join or avoid others. Moreover, within a given species, the decision to join or avoid others can be affected markedly by the physiological and informational state of the individual and by contextual response thresholds to resource availability. Decisions that benefit the individual may or may not affect the group in terms of total reproductive output.

Host odor and visual stimulus interaction during intratree host finding behavior ofRhagoletis pomonella flies.

Responses ofRhagoletis pomonella (Walsh) (Diptera: Tephritidae) flies to host fruit visual stimuli (apples or models of apples) and chemical stimuli (synthetic apple volatile blend) were studied in semidwarf field-caged apple trees. Three different fruit or model densities (1, 4, or 16 fruit or models/ tree) and two odor release rates [ca. 0.7µg/hr (close to the natural release rate of a ripe apple) and ca. 500µg/hr (amount of odor released by commercially sold apple maggot traps)] were tested. Individually released flies were followed as they moved within a tree for a maximum of 20 min. We recorded three-dimensional search paths followed by foraging flies and computed such variables as total relative distance traveled before alighting on a fruit or model, track length between individual alightment sites, and directness of flight to fruits or models. Effect of odor on propensity to alight on fruit or models and host-searching behavior prior to alighting on fruit or on models varied according to fruit or model color and density. If the fruit visual stimulus was strong (e.g., red color), odor did not increase the probability of finding fruit or fruit models. As the visual stimulus became progressively weaker (red to green to clear), odor (irrespective of concentration) appeared to aid flies during the fruit-finding process. As density of fruit or models increased, the probability of flies finding a fruit or model also increased (e.g., 50% of flies found a red fruit model at 1 model/tree while 90% found a red model at 16 models/tree; 4% of flies found a clear model with odor at 1 model/tree while 35% found a clear model with odor at 16 models/tree). Findings reported elsewhere indicate thatR. pomonella flies are able to discover a point source of odor (an odor-bearing tree in a patch of trees) by flying upwind (in the tree patch) in response to intermittent exposure to odor. Findings here indicate that after arrival on a host tree (point source), flies discover individual apparent and abundant host fruit on the basis of vision. If fruit are less apparent or scarce, odor appears to interact with vision during the fruit-finding process.

The effect of short-range host odor stimuli on host fruit finding and feeding behavior of plum curculio adults (Coleoptera: Curculionidae).

In laboratory assays, we investigated responses of female plum curculios (PCs),Conotrachelus nenuphar (Herbst), to host and nonhost fruit or leaf odor when PCs were crawling on experimental tree branchlets or twigs. In choice tests where test specimens were hung from the ends of a wooden crosspiece, PCs made significantly more visits to host plum fruit than to plum leaves, nonhost tomato fruit, wax models of plum fruit, or blanks (wire). In similar tests, PCs made significantly more visits to plum leaves compared to nonhost maple leaves or to blanks. PCs in test chambers that contained host or nonhost odor were significantly more prone to feed on wax plum models in the presence of odor from host fruit or host leaves compared to odor from nonhost fruit or leaves or a water blank. In choice tests offering alternating cluster types on an apple branchlet, PCs visited leaf clusters bearing a host apple fruit more than leaf clusters without a fruit. In tests to assay the distance at which PCs can detect an individual host fruit, PCs crawled from the central stem of an apple branchlet onto a side stem significantly more often when an apple fruit on a side stem was hung 2 cm from the central stem compared to 4 or 8 cm away. Our combined results suggest that PCs use host fruit odor to locate host fruit at close range.

Attraction of apple maggot flies, Rhagoletis pomonella (Diptera: tephritidae) of different physiological states to odour-baited traps in the presence and absence of food.

Adults of apple maggot fly Rhagoletis pomonella (Walsh) of differing physiological states were marked and released in blocks of apple trees ringed by sticky red spheres. Spheres were either unbaited, baited with butyl hexanoate (synthetic host fruit odour) or baited with both butyl hexanoate and ammonium carbonate (synthetic food odour). All trap and lure treatments were compared in the presence or absence of food (bird faeces) in the blocks. Simultaneously, the response of wild immigrant flies to treatments was measured and wild females were dissected to determine state of ovary development. Large proportions (25-40%) of released mature male and female R. pomonella were recovered in blocks having traps baited with butyl hexanoate. Ammonium carbonate did not enhance trap captures and presence of food had little effect on response to synthetic odours by mature R. pomonella. Immature flies of each sex responded weakly to traps and to both types of synthetic lures and may have been arrested in blocks having food. Wild flies of both sexes exhibited a response pattern very similar to mature released flies, regardless of eggload (in the case of wild females). Results indicate that wild R. pomonella immigrating into apple orchards are primarily mature, and not hungry for protein. Behavioural control strategies are discussed in that context.

Mite predator responses to prey and predator-emitted stimuli.

We found that the searching behavior of two acarine predators,Amblyseius fallacis andPhytoseiulus macropilis, for prey,Tetranychus urticae, is affected by the following stimuli: (1) prey silk and associated feces, whose combined physical and chemical properties elicit reduction in the rate of predator movements and longer halts; (2) kairomone extracted from prey silk and associated feces, which, upon contact, elicits frequent predator return to prey-inhabited locales; and (3) predator-emitted marking pheromone, which elicits shorter duration of search in presearched prey locales. We also found that treatment of filter paper with prey kairomone or silk enhanced predator location of prey eggs, leading us to speculate that application of synthetic prey kairomone could be useful in pest management programs.

Residual activity of oviposition-deterring pheromone inRhagoletis pomonella (Diptera: Tephritidae) and female response to infested fruit.

Under dry conditions, oviposition-deterring pheromone (ODP) of the apple maggot fly (Rhagoletis pomonella Walsh) on host fruit proved deterrent to ovipositing females for at least three weeks, with a half-life of ca. 11 days. There was no difference in decline of residual activity under lab vs. field conditions or between fly-deposited ODP vs. an application of a water extract of ODP. A decline in pheromone activity resulted from exposure to both natural and simulated rainfall. For natural rainfall, greatest losses (50- 61%) in activity resulted from high-intensity rains, with substantially less activity lost (13-35%) following light or moderate intensity rains.R. pomonella females discriminated against fruit without ODP but with conspecific larvae. Discrimination against infested fruit was manifested within fewer days following infestation of small fruit (9 mm diam.) as compared to larger fruit (15 mm diam.).

Phytochemical basis of learning inRhagoletis pomonella and other herbivorous insects.

Examples of phytochemically-based learning of host preference in herbivorous insects are reviewed in the context of traditionally important issues: the number and kinds of chemicals involved; which sensory modalities are affected; whether peripheral or central nervous processing is altered; and whether learning is associative or not. A fifth issue addressed here- whether experience enhances a feeding or ovipositing insect's propensity to accept familiar chemical stimuli or to reject novel chemical stimuli-has been ignored in previous studies. Following the review, evidence is presented indicating that female apple maggot flies (Ragoletis pomonella) learn to reject both novel physical and novel chemical stimuli.

Distance of response to host tree models by female apple maggot flies,Rhagoletis pomonella (Walsh) (Diptera: Tephritidae): Interaction of visual and olfactory stimuli.

Mature female apple maggot flies,Rhagoletis pomonella (Walsh), were released individually onto a single potted, fruitless hawthorne tree in the center of an open field. The tree was surrounded by four 1-m(2) plywood host tree models painted green or white, with or without synthetic host fruit odor (butyl hexanoate), and placed at one of several distances from the release tree. Each fly was permitted to forage freely on the release tree for up to 1 hr, or until it left the tree. Flies left the tree significantly sooner when green models with host fruit were present at 0.5, 1.5, or 2.5 m distance from the release tree than when these models were placed at a greater distance (4.5 m) from the release tree or when no models were present. Flies responded detectably to 1-m(2) models without odor up to a maximum distance of 1.5 m. These results suggest that female apple maggot flies did not detect green 1-m(2) models with odor 4.5 m away or models without odor 2.5 m or more away. Flies responded to white models with and without odor to a much lesser extent, both in terms of response distance and flight to and alightment upon models. Increasing model size to 2 m(2) increased the distance to 2.5 m at which flies responded to green models without odor. Decreasing model size to 0.5 m(2) reduced fly responsiveness to green or white models. The presence of host fruit odor alone, without the visual stimulus of a green model, did not influence residence time on the release tree.

Factors influencing release of hostmarking pheromone byRhagoletis pomonella flies.

The effect of fly or fruit treatments on quality and/or quantity of host-marking pheromone (HMP) trail substance released by apple maggot flies (Rhagoletis pomonella) following oviposition was evaluated. Among flies, considerable variation existed in the amount of HMP substance deposited, but overall, the amount of substance released on successively offered fruit (over a day or a week) did not change appreciably. Fly diet did not influence pheromone activity. Older flies (28 days) or smaller flies released less or less active HMP trail substance than younger flies (14 days) or larger flies. Females deposited a similar amount of trail substance on large (18-19 mm diam.) or HMP-marked fruit as on small (12-13 mm) or unmarked fruit. Starvation reduced the amount of measurable trail substance deposited but resulted in a more active HMP deposition. Discrepancy between trail measurement and behavioral bioassay results for the starvation treatment indicated that trail measurement results may be misleading under conditions that reduce gut contents of the fly.

Bioassay approaches to assessing behavioral responses of plum curculio adults (Coleoptera: Curculionidae) to host fruit odor.

We evaluated several approaches to developing a simple, sensitive, and reliable laboratory bioassay of responses of overwintered adult plum curculios (PCs),Conotrachelus nenuphar (Herbst), to host fruit odor or its attractive components. A high proportion of assayed PCs responded positively to odor of wild plums under no-choice, moving-air conditions in a wind tunnel and under dual-choice, still-air conditions in enclosed Petri dishes. Positive response to controls lacking host odor, however, was much greater in the wind tunnel, arguing in favor of bioassays under dual-choice conditions in still air to provide greater PC discrimination. Response to host odor (from wild plums or hexane extract of wild plums or Liberty apples) in Petri dish bioassay chambers proved greatest: (1) during the scotophase of PCs under total dark or dim red light conditions, (2) when Petri dishes were completely enclosed, (3) when PCs were starved for 24 or 48 hr, and (4) when PCs were tested within seven weeks after apple tree petal fall. Neither the sex of a PC nor the direction in which a PC was obliged to move to find the source of host odor (upward through a port in the Petri dish lid or downward through a port in the base) had a substantial effect on level of response to host odor or discrimination of host odor from a nonodorous control. We conclude that an enclosed Petri dish bioassay chamber of the type described here should be a valuable asset in the process of chemically identifying components of host fruit odor attractive to PCs.