Semiochemical-Mediated Oviposition Avoidance by Female House Flies, <Emphasis Type="Italic">Musca domestica</Emphasis>, on Animal Feces Colonized with Harmful Fungi

House flies, Musca domestica, utilize ephemeral resources such as animal feces for oviposition and development of larval offspring, but they face competition with fungi that colonize the same resource. We predicted that house flies avoid oviposition on feces well-colonized with fungi, thereby reducing fungal competition for larval offspring. Working with fungal isolates from chicken feces, we have previously shown that prior establishment of Phoma spp., Fusarium spp., or Rhizopus spp. on feces significantly reduced oviposition by house flies. Here, we report that, in the headspace volatiles of these three fungal genera, five compounds (dimethyl trisulfide, an unknown, 2-phenylethanol, citronellal, norphytone) elicit responses from house fly antennae. In behavioral bioassays, dimethyl trisulfide and 2-phenylethanol significantly reduced oviposition by house flies. We conclude that fungus-derived volatiles serve as semiochemical cues that help house flies avoid resources colonized with fungal competitors for the development of larval offspring.     

Quantity Does Matter: How Feces Are Used for Host Stage Selection by Granary Weevil Parasitoid Lariophagus distinguendus

The present paper examines the mechanisms of host stage selection in Lariophagus distinguendus, a larval parasitoid of the granary weevil Sitophilus granarius endophytic in wheat grain. The amount of host feces accumulating in infested grains was on average, 0.2, 0.3, 1.5, and 4 mg for first, second, third, and fourth instars, respectively. A maximum of 9 mg feces was found in grains with fourth instars. In bioassays with female parasitoids, only a little drumming and drilling behavior was observed on grain models treated with feces extract equivalent to 0.2 mg and 0.4 mg feces. Most drumming and drilling occurred on models with extracts equivalent to 4 and 8 mg feces. These results correspond to literature data demonstrating that older host larvae (fourth instars) are preferred for oviposition. Thus, host stage selection in L. distinguendus is apparently achieved by assessing the quantity of feces present in infested grains.     

Aggregation pheromones ofDrosophila immigrans,D. phalerata,andD. subobscura

Aggregation pheromones ofDrosophila immigrans, D. phalerata andD. subobscura were demonstrated by testing attraction of adult flies to hexane extracts of the flies in a windtunnel bioassay. Extracts of adult males of all species attracted conspecific males and females. However,D. subobscura flies were attracted only when the extract (cVA) in the extracts of adult maleD. immigrans andD. phalerata. Both species were attracted to synthetic cVA. Male and femaleD. phalerata. Both species were attracted to synthetic cVA. Male and femaleD. subobscura produced 5,9-pentacosadiene, 5-pentacosene, 2-methylhexacosene and 5,9-heptacosadiene, while only maleD. subobscura produced (Z)-5-tricosene and minor amounts of cVA.     

Host marking pheromone ofRhagoletis cerasi: Foraging behavior in response to synthetic pheromonal isomers

We studied the foraging behavior ofRhagoletis cerasi females in trees treated with synthetic cherry fruit fly host marking pheromone (HMP) under seminatural conditions (potted trees enclosed in a screen cage). Results show that synthetic HMP (particularly the 8RS-@#@ 15R isomer configuration (racemic mixture)) was highly effective in eliciting behavioral responses similar to those reported in studies using natural HMP. Flies exposed to synthetic pheromone exhibited short tree residence times (i.e., emigrated faster), increased flight frequency rates (measured as number of alightings per/minute), higher irritation indices while on a tree or a fruit, and oviposited fewer eggs per fruit visit than flies exposed to clean trees and fruit (not treated with synthetic HMP). Furthermore, we provide evidence showing that when flies were continuously exposed to an HMP-saturated environment, they exhibited an increased tendency to lay eggs in marked fruit.     

Host-Plant Chemistry Influences Oviposition Choice of the Spicebush Swallowtail Butterfly

The spicebush swallowtail, Papilio troilus (L.), lays its eggs on plants in the family Lauraceae. Sassafras [Sassafras albidum (Nutt.) Nees], spicebush [Lindera benzoin (L.) Spreng.], redbay (Persea borbonia (L.)] and camphortree [Cinnamomum camphora (Nees) Eberm.] are four of its known host plants. In one-choice tests, free-flying spicebush swallowtail females laid eggs on chemical extracts of the leaves of each of these four hosts. In two-choice experiments, females always preferred to oviposit on an extract of sassafras compared to extracts of the other three hosts. It was shown for spicebush extract that this response was not due to oviposition experience. Previously we had identified one of the host plant chemicals acting as an oviposition stimulant in sassafras extract as 3-caffeoyl-muco-quinic acid (3-CmQA). Extracts of the other three hosts did not contain this compound. The addition of 3-CmQA alone to spicebush extract did not increase oviposition activity. It did, however, increase discrimination between hosts and nonhosts. When a fraction of sassafras extract containing 3-CmQA and other synergistic stimulants was added to spicebush extract, preference for sassafras extract was no longer recorded. These results show existing differences in oviposition chemistry among host plants of the spicebush swallowtail and how these differences can influence oviposition choice in bioassay experiments.     

Deterrence of repeated oviposition by fruit-marking pheromone inCeratitis capitata (Diptera: Tephritidae)

During ovipositor dragging on the fruit surface following egg laying in hawthorne fruit,Ceratitis capitata (Wiedemann) females deposit an unidentified pheromone that deters repeated oviposition attempts in that fruit. The pheromone proved water soluble and, when collected and sprayed in aqueous solution onto uninfested fruits in laboratory cages, effectively deterred boring attempts byC. capitata females of wild origin for at least 6 days (termination of test). A laboratory population ofC. capitata cultured on artificial media for more than 200 generations deposited pheromone that proved equally as deterrent to wild fly oviposition as pheromone from wild flies. However, lab fly oviposition was not effectively deterred by the presence of pheromone. The ecological significance of the pheromone is discussed.     

House fly oviposition inhibition by larvae ofHermetia illucens,the black soldier fly

Wild populations of house flies were inhibited from ovipositing into poultry manure containing larvae of the black soldier fly,Hermetia illucens (L.). A laboratory strain of house fly responded differently, readily ovipositing into manure with lower densities of soldier fly larvae, but avoiding the higher densities tested. The amount of timeH. illucens larvae occupy the manure prior to an oviposition test influences ovipositional responses of house flies. Manure conditioned byH. illucens larvae for 4–5 days did not significantly inhibit house fly oviposition. We suggest that some type of interspecific chemical communication (allomone) is present.Diptera: Stratiomyidae.Portion of an MS thesis by S.W.B.Research partially supported by State and Hatch funds.     

d-(+)-Pinitol,an oviposition stimulant for the pipevine swallowtail butterfly,Battus philenor

Oviposition by females of the pipevine swallowtail butterfly,Battus philenor, was stimulated by contact with alcoholic extracts of host foliage.d-(+)-Pinitol was isolated and identified from leaf material of one host species,Aristolochia macrophylla (Aristolochiaceae). In combination with chloroform-soluble components of host leaf material, this compound was comparable to the parent extract in stimulating oviposition.     

Identification of Host Fruit Volatiles from Flowering Dogwood (Cornus florida) Attractive to Dogwood-Origin Rhagoletis pomonella Flies

Solid-phase microextraction and gas chromatography coupled with electroantennographic detection were used to identify volatiles from fruit of flowering dogwood, Cornus florida, as key attractants for Rhagoletis pomonella flies originating from dogwood fruit. A six-component blend containing ethyl acetate (54.9%), 3-methylbutan-1-ol (27.5%), isoamyl acetate (0.9%), dimethyl trisulfide (1.9%), 1-octen-3-ol (9.1%), and -caryophyllene (5.8%) was identified from flowering dogwood fruit that gave consistent EAD activity. In a flight tunnel assay there was no significant difference in the response of individual dogwood flies exhibiting upwind anemotactic flight to volatile extracts from dogwood fruit and the six-component synthetic mixture. Dogwood flies also displayed significantly greater levels of upwind flight to sources with the dogwood volatile blend than with previously identified volatile blends from domestic apple or hawthorn fruit. Selected subtraction assays showed that the three-component mixture of 3-methylbutan-1-ol, 1-octen-3-ol, and -caryophyllene elicited levels of upwind flight to the source equivalent to the six-component mixture. Our study adds to previous ones showing that populations of Rhagoletis pomonella flies infesting apple, hawthorn, and flowering dogwood fruit are attracted to unique mixtures of fruit volatiles, supporting the hypothesis that host fruit odors could be key traits in sympatric host shifts and establishing host fidelity within members of the Rhagoletis pomonella species complex.     

Response to walnut olfactory and visual cues by the parasitic wasp Diachasmimorpha juglandis

Diachasmimorpha juglandis is a specialist parasitoid attacking fly larvae in the genus Rhagoletis that feed exclusively on walnut fruit husks. In a free-foraging assay comparing response to uninfested, infested, and mechanically damaged fruits, we first determined that D. juglandis use host feeding damage on the fruit as a cue for host presence. In another free-foraging assay that used artificial walnut models and wind tunnel experiments, D. juglandis distinguished infested from uninfested fruits by using either olfactory or visual cues separately. However, the response rate of wasps in the wind tunnel was raised considerably when visual cues were also available. We analyzed the volatile compounds emitted by cohorts of uninfested, mechanically damaged, and infested fruits 1, 3, 5, 8, and 10 days after oviposition by flies into the infested fruits. Total volatile emissions did not differ significantly among treatments, but quantitative changes in volatiles distinguished infested fruits from uninfested and damaged fruits. The fact that parasitoids did not distinguish between infested and damaged fruits in assays where damage was visible indicates that they rely on visual cues when those are available.     

Volatiles from Duck Feces Attractive to Mexican Fruit Fly

Aqueous extracts of three types of bird feces were attractive to laboratory-colony Mexican fruit flies in laboratory bioassays. Extracts of black-bellied whistling duck feces were chosen for further analysis. Duck feces extract was attractive to both sugar-fed and sugar-starved flies but was more attractive to sugar-fed flies. Protein deprivation of flies had no effect on their response. Duck feces incubated in water for 24 hr were more attractive than those incubated for longer periods. Duck feces extract was attractive at pH 5–9, but was most attractive at pH 9, indicating that more than one class of chemicals contributed to attractiveness of the feces. Major components of headspace of duck feces extracts collected by solid-phase microextraction and identified by GC-MS, GC-FID, and GC-FTD included ethanol, propanol, phenol, ammonia, low-molecular weight amines, and pyrazines. In general, chemicals containing nitrogen proved most attractive to sugar-fed flies. However, dimethylamine and 1-pyrroline were nearly equally attractive to sugar-fed and sugar-deprived flies. Two chemicals without nitrogen were attractive to sugar-deprived flies; none were attractive to sugar-fed flies. A synthetic mixture was prepared containing ammonia, methylamine, dimethylamine, trimethylamine, 1-pyrroline, phenol, and 2-ethylhexanol in amounts found in odor of duck feces extract. The synthetic mixture was 96% as attractive as duck feces extract to sugar-fed flies and 80% as attractive to sugar-deprived flies.     

Oviposition stimulants for the tropical swallowtail butterfly,Papilio polytes,feeding on a rutaceous plant,Toddalia asiatica

In nature, Papilio polytes utilizes a limited range of rutaceous plants as hosts. We isolated and identified oviposition stimulants for the butterfly from the foliage of its primary host plant Toddalia asiatica. Females readily deposited eggs in response to a methanolic extract of the plant. Partition of the extract with organic solvents revealed that chemicals responsible for eliciting egg-laying resided in a water-soluble fraction. Further bioassay-guided fractionation of the active fraction by column chromatography, preparative TLC, and HPLC led to the isolation of two oviposition stimulants. One was isolated from an amphoteric fraction and identified as trans-4-hydroxy-N-methyl-L-proline [(–)-(2S,4R)-4-hydroxy-1-methyl pyrrolidine-2-carboxylic acid; HMP]. The other, isolated from an acidic fraction, was identified as 2-C-methyl-D-erythronic acid [(–)-(2R,3R)-2-methyl-2,3,4-trihydroxybutanoic acid; MEA]. HMP alone evoked significant oviposition-stimulatory activity, although this was much lower than that of the original water-soluble fraction. MEA, on the other hand, alone did not elicit positive responses from females. However, HMP, when assayed in combination with MEA, markedly enhanced the female response, and the mixture was as active in stimulating oviposition as were the original water-soluble fraction and the plant foliage. We conclude that HMP is a substance crucial for host recognition by females, while MEA is a synergistic stimulant involved in host recognition and/or preference.     

Plant response to eggs vs. Host marking pheromone as factors inhibiting oviposition byPieris brassicae

Pieris brassicae L. butterflies secrete miriamides onto their eggs. These avenanthramide alkaloids are strong oviposition deterrents when sprayed onto a cabbage leaf. However, these compounds could not be detected in cabbage leaves from which egg batches had been removed two days after deposition and that still showed oviposition deterrency. It was concluded that the miriamides were not directly responsible for the avoidance by females of occupied leaves while searching for an oviposition site. Evidence was obtained that cabbage leaves themselves produce oviposition deterrents in response to egg batches. Fractions containing potent oviposition deterrents could be isolated from surface extracts of leaves from which previously laid egg batches had been removed. The term host marking pheromone that was used previously is not applicable in this case.     

Molecular basis ofMorinda citrifolia (L.): Toxicity on drosophila

The ripe fruit ofMorinda citrifolia, host plant forDrosophila sechellia is highly toxic for three closely related species (D. melanogaster, D. simulans, andD. mauritiana). Green and rotten fruits are not toxic for all species tested. Short fatty acids were found to be present in large quantities in the extract of the ripe fruit. The most abundant (octanoic acid) was tested pure for its toxicity in a dose-dependent manner;D. sechellia is five to six times more resistant thanD. melanogaster to octanoic acid. Octanoic acid alone seems to be sufficient to explain the toxic effect of the pulp. It is less abundant in the rotten fruit and absent in the green fruit.     

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-m² 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-m² 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-m² 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 m² increased the distance to 2.5 m at which flies responded to green models without odor. Decreasing model size to 0.5 m² 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.     

Differences among sibling speciesRhagoletis mendax andR. pomonella (diptera: ephritidae) in their antennal sensitivity to host fruit compounds

Prior electrophoretic and morphological studies have identified two closely related, economically important tephritid flies,R. mendax (Curran) andR. pomonella (Walsh), which infest the fruits of ericaceous and rosaceous plants, respectively. Further studies also have shown consistent differences among these species in their ovipositional preferences for apples and highbush blueberries and have determined that their ovipositional behavior is elicited by extracts obtained from these fruits. In this paper we report the results of an experiment that tested whether these species show distinct electroantennogram (EAG) responses to a large array of compounds present in gas chromatograph-fractionated pentane extracts of apples and highbush blueberries.R. mendax andR. pomonella flies were found to have significant differences in their antennal sensitivity to 11 blueberry and nine apple extract peaks, which correspond to 24.4% of all blueberry and 25.0% of all apple peaks that elicited a measurable EAG response from either species. Interspecific differences in peripheral sensitivity were more pronounced for blueberry than apple extract;R. pomonella flies were most sensitive to blueberry compounds with low retention times, whereasR. mendax flies responded to blueberry compounds with a broader range of retention times. Both species were most sensitive to apple peaks with high retention times. The retention times of most apple and blueberry peaks that elicited EAG responses fromR. mendax andR. pomonella flies were different from the retention times of seven attractant fruit esters that were previously identified by Fein et al. (1982). The identification of these unknown apple and blueberry compounds could lead to the discovery of new chemical cues that mediate the host-plant preferences of these sibling species.     

Oviposition in Delia platura (Diptera, Anthomyiidae): The Role of Volatile and Contact Cues of Bean

The choice of a suitable oviposition site by female insects is essential for survival of their progeny. Both olfactory and contact cues of the oviposition site may mediate this choice. The polyphagous Delia platura (Diptera: Anthomyiidae), a severe agricultural pest of numerous crops, lays eggs in the soil close to germinating seeds. Maggots feed upon the cotyledons. Only little is known about the cues guiding oviposition behavior. In this study, the effects of both olfactory and contact cues of beans (Phaseolus vulgaris) on oviposition of D. platura females were tested. Egg deposition on germinated beans was preferred to egg deposition on ungerminated beans or on beans in different postgerminating developmental stages. Olfactory cues of germinating beans alone stimulated female flies to lay eggs. Additional contact cues of germinating beans seemed to enhance the response, but the difference was not significant. Surface extracts of germinating beans sprayed on surrogate beans showed that both polar and nonpolar substances stimulated oviposition of D. platura flies. Gas chromatography-electroantennographic detection recordings of head space samples of germinating beans showed positive response of females to different compounds. We conclude that olfaction plays a major role when D. platura females are searching for oviposition sites. Volatile compounds released from germinating beans such as 4-hydroxy-4-methyl-2-pentanone, 1-hepten-3-one, 1-octen-3-ol, and 3-octanone should be considered as key compounds that mediate oviposition behavior. The use of different sensory modalities by closely related species of Delia is discussed.     

Evidence for a Host-Marking Pheromone in White Spruce Cone Fly, Strobilomyia neanthracina

Cone flies (Strobilomyia spp.) lay eggs on coniferous cones, and larvae must complete development within the cone on which the eggs are laid. Previous field surveys showed that egg distributions of several species are uniform, suggesting that females avoid ovipositing on cones with conspecific eggs or larvae. In both the field and laboratory, S. neanthracina females walked around cones following oviposition, touching their mouthparts to the cone as they walked. In laboratory bioassays, where females were presented with a cone on which they or another female had oviposited and touched with mouthparts, or one without an egg, females laid preferentially on cones without eggs. However, females laid randomly when presented with a cone on which a female had oviposited but been prevented from touching with her mouthparts following oviposition or another cone without an egg. This indicates that females deposit a host-marking pheromone with their mouthparts following oviposition and that this deters further oviposition on marked hosts. Laboratory and field bioassays indicated that the host-marking pheromone is located in the head and thorax of female flies.     

Effects on behavior ofApanteles melanoscelus females caused by modifications in extraction,storage, and presentation of Gypsy moth silk kairomone

Choice experiments were performed to investigate details of femaleApanteles melanoscelus (Ratzeburg) behavior when exposed to gypsy moth silk kairomone [Lymantria dispar (L.)] and to host larvae when kairomone is present. Female parasites only responded to the kairomone when it had been placed on thin strands such as cotton fibers. Both gypsy moth silk and silk glands contain the same or similar kairomones. Silk gland extracts were more active than head, alimentary canal, or hemolymph extracts of host larvae. Female responses decreased when low concentrations of silk gland extract or small numbers of treated fibers were presented to them. Silk gland extract was stable when frozen for 2 weeks, heated to 100 °C for 0.5 hr, freeze dried, or treated with 95% ethanol. The active component was nondialyzable. Silk deposited on the substrate increased host contacts and oviposition attempts, more so if wider areas were covered with silk, and even if the areas having silk were separated from the host. A theory of host selection inA. melanoscelus is proposed.Hymenoptera, Braconidae. Lymantria dispar L. (Lepidoptera: Lymantriidae).     

Host egg kairomones essential for egg-larval parasitoid,Ascogaster reticulatus watanabe (Hymenoptera: Braconidae)

An artificial egg with a Parafilm membrane was devised for the oviposition ofAscogaster reticulatus Watanabe (Hymenoptera: Braconidae), an egg-larval parasitoid of the smaller tea tortrix,Adoxophyes sp. (Lepidoptera: Tortricidae). Both external and internal kairomones were essential. The external kairomone, needed for host location and acceptance, was extracted with 70% ethanol, and the internal kairomone, needed for oviposition, was extracted with water. Female parasitoids responded to the external kairomone and oviposited through the membrane into the artificial egg when the supernatant of host egg-mass homogenate was inside, whereas they did not when water or saline solutions were inside. Thus an internal kairomone is responsible for the oviposition in the host egg. The internal kairomone apparently was not specific for the host egg mass because oviposition activity was found not only in egg, larval, and pupal stages of the host, but also in larvae of other species of Lepidoptera and Coleoptera.