Trimedlure affects mating success and mate attraction in male Mediterranean fruit flies

Male Mediterranean fruit flies, Ceratitis capitata (Wiedemann), are attracted to the synthetic compound trimedlure. Despite the common use of trimedlure in control programs, the underlying basis of male attraction remains unknown. In a series of laboratory trials, we compared the mating success of (treated) males exposed to trimedlure and (control) males not given access to trimedlure. When tested immediately after exposure, treated males mated more frequently than control males. This mating advantage was short-lived, however, and males tested as little as 24 h after exposure had no advantage over control males. When tested immediately after exposure, treated males exhibited higher levels of pheromone calling than control males. Consequently, more females were attracted to treated males than to control males in field tests.     

Behavioural switch of female Mediterranean fruit fly, Ceratitis capitata: mating and oviposition activity in outdoor field cages in Hawaii

1 Laboratory-reared normal, and wild female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), were assayed in outdoor field cages to assess the impact of a mating-induced behavioural switch on mating and subsequent oviposition activity. 2 Virgin females preferred interactions with males leading to mating over attraction to, and oviposition in, artificial yellow spheres containing guava odour or green apples hung in a guava tree. Laboratory-reared females previously mated with either laboratory-reared normal males or laboratory-reared irradiated (sterile) males showed little interest in remating with males and instead, were much more likely to be found arrested on artificial and real fruit and ovipositing. Oviposition on artificial fruit was five times greater by females that had mated with either normal or irradiated males than by virgin females. Wild females showed similar qualitative changes in the mating-induced behavioural switch; however, oviposition activity was significantly less than for laboratory-reared females. 3 These results confirm that mating has a profound effect on the behaviour of female Mediterranean fruit flies and that irradiated males are functionally equal with normal males (lab-reared or wild) in their ability to alter female behaviour. These results are discussed in the context of the sterile insect technique for control of Mediterranean fruit flies in the field.     

What Do Mexican Fruit Flies Learn When They Experience Fruit?

Mexican fruit flies learn fruit characteristics that enable them to distinguish familiar fruits from novel fruits. We investigated whether mature Mexican fruit flies learn fruit color, size or odor. We found no evidence that female flies learn fruit color or size after experience with host fruit, including oviposition. However, green fruit and fruit models were more attractive than yellow and red fruit and fruit models regardless of previous experience. Females with grapefruit experience were more attracted to fruit models with extract of either grapefruit peel or pulp, than to models without extract. Females with no experience with grapefruit were not attracted to models treated with grapefruit extract. These results indicate that females learned fruit odor during exposure to grapefruit.     

Colonization of a hybrid strain to restore male Anastrepha ludens (Diptera: Tephritidae) mating competitiveness for sterile insect technique programs

To restore male mating competitiveness of Mexican fruit flies, Anastrepha ludens (Loew) (Diptera: Tephritidae), reared for sterile insect releases by the Mexican Fruit fly Eradication Campaign, two strain replacement techniques were evaluated. Field cage male competitiveness tests revealed that laboratory males of the Metapa strain mated 3 times less often with wild females than field-collected wild males. A strain developed from the cross of wild males and laboratory females (hybrid strain) was similar to a strain developed from the cross of laboratory males and females (laboratory strain) in that its females produced similar amounts of eggs and the eggs displayed similar levels of hatch and egg-to-pupa transformation in artificial diet. By contrast, a strain developed from the cross of wild males and females (wild strain), forced into artificial rearing, experienced a series of bottlenecks involving reduced egg laying and extremely poor development in diet. The male F1 progeny of the hybrid strain and field-collected wild males outcompeted Fl laboratory males in field cage tests for matings with field-collected wild females. In conclusion, we found that strains developed from the cross of wild males and laboratory females allowed us to restore male mating competitiveness of F1 Mexican fruit flies without compromising mass-rearing production.     

Mass rearing history negatively affects mating success of male Anastrepha ludens (Diptera: Tephritidae) reared for sterile insect technique programs

Mating competitiveness and sterility induction into cohorts of wild Anastrepha ludens (Loew) (Diptera: Tephritidae) was compared among wild and laboratory flies reared for use in the sterile insect technique Mexican program. Laboratory flies stemming from an 11-yr-old bisexual strain were either not irradiated, irradiated at 3 krad (low dose), or irradiated at 8 krad. In 30 by 30 by 30-cm Plexiglas cages, where a cohort of laboratory flies (male and female) irradiated at different doses (0, 3, and 8 krad) was introduced with a cohort of wild flies, males and females of each type mated randomly among themselves. Compared with nonirradiated laboratory and wild males, irradiated males, irrespective of dose (3 or 8 krad), induced shorter refractory periods and greater mating frequency in wild females. Nevertheless, laboratory flies irradiated at a low dose induced greater sterility into cohorts of wild flies than laboratory flies irradiated at a high dose. In a 3 by 3 by 3-m walk-in cage, wild males gained significantly more matings with wild females than nonirradiated and irradiated laboratory males a finding that revealed a strong effect of strain on mating performance. Mating incompatibility of the laboratory strain might have obscured the effect of reduced irradiation doses on male mating performance in the walk-in cage. Our results highlight an urgent need to replace the A. ludens strain currently used by the Mexican fruit fly eradication campaign and at least suggest that reducing irradiation doses result in an increase in sterility induction in wild populations.     

Attraction of the cabbage looper to host plants and host plant odor in the laboratory

In a laboratory flight tunnel, mated female, unmated female, and male adult cabbage loopers, Trichoplusia ni (Hübner), exhibited chemically mediated anemotaxis (attraction) in response to intact potted cabbage plants (Brassica oleracea L.), leading to contact with the plant. Similar attraction responses were also observed by mated females to potted soybean (Glycine max (L.)), tomato (Lycopersicon esculentum Miller), and celery (Apium graveolens L.) plants in noncompetitive comparisons. Mated females, unmated females, and males flew to cabbage plants throughout the scotophase. Response rates for mated females were higher than for unmated females and males. Mated female cabbage loopers were attracted by odors of cabbage, soybean, tomato, or celery piped into a flight tunnel from single plants held in glass jars and not to odors of the non-host plant Setcreasea purpurea, or to humidified air. They were also attracted to water washings of cabbage at dosages of 0.4 to 2.0 gram equivalents, when presented on cotton dental wicks in the flight tunnel.     

Reduction in the attraction of males to females after mating in the rice leaf bug Trigonotylus caelestialium

The influence of mating on the extent to which males are attracted to females in Trigonotylus caelestialium (Heteroptera: Miridae) was examined. No differences in attraction of males to mated and virgin females were observed within 3–5 h of mating, but males became less attracted to females 1 to 2 days after the first mating. The difference in male attraction to mated vs virgin females disappeared at 4 days after mating. These results indicate that reduced attraction of males to mated females occurs after a certain time interval, and persists for a few days. Furthermore, males were less attracted to females that had mated with virgin vs recently mated males, i.e. males that had just mated with another female at 1 and 2 days after mating. The ejaculate expenditure of recently mated males was less than that of virgin males. Hence, the amount of male ejaculate transferred to females during mating, rather than the act of mating, might influence the attraction of males to females. The results demonstrate that mating reduces the attraction of males to females in T. caelestialium on the basis of direct observation of male behavior.     

Flight Tunnel Responses of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) to Olfactory and Visual Stimuli

Cohorts of mass-reared, naive Diachasmimorpha longicaudata, parasitoids of tephritid fruit flies, were released in a laminar airflow wind tunnel to study their responses to visual and olfactory stimuli associated with their host habitat. Parasitoids were five times more likely to land on yellow plastic spheres emitting the odor of ripe, guava fruit (Psidium guajava L.) than to spheres emitting clean air. The rate of landing was not modified by the presence of green artificial leaves adjacent to the spheres in the tunnel or by the inclusion of green leaf volatiles emanating with the guava odors. However, hovering activity was twice as frequent around spheres adjacent to artificial leaves than around isolated spheres. Oviposition activity on spheres emitting guava odor was not affected by the presence of artificial leaves nor by green leaf volatiles. This suggests that inexperienced D. longicaudata may be instinctively attracted to foliage and to fruit odor but that landing (arrestment) and oviposition are influenced more by odor than by the appearance of fruit or foliage. D. longicaudata are not instinctively attracted to larvae of Bactrocera dorsalis in the absence of host-habitat stimuli. More wasp activity occurred around oviposition units containing larvae if the odor of ripe/overripe guava was present. Successful wasp reproduction occurred only in units with guava odor.     

Electroantennogram response and attraction of Anastrepha suspensa to volatiles of various sugar sources and aged sugar solutions

With the aim of finding new, sugar‐based volatile attractants for economically important tephritid fruit fly species, we used electroantennography (EAG) to quantitate olfactory responses of female Caribbean fruit fly, Anastrepha suspensa (Loew) (Diptera: Tephritidae), to volatiles of six sugar sources (refined white and brown cane sugar, coconut sugar, date sugar, date jaggery, and cane panela). Laboratory‐strain and wild flies, both sexually immature and mature, were tested for EAG responses to the volatiles of dry crystallized sugar sources and 10% (wt/vol) aqueous solutions that had aged in the laboratory for 0–7 days. In general, wild flies exhibited higher EAG responses than laboratory flies, and immature females responded more strongly than mature females. With the exception of date jaggery and cane panela, volatiles of dry sugar sources and 0‐ and 1‐day‐old solutions elicited lower EAG responses than any of the aged solutions. Most solution volatiles elicited the strongest EAG response after 2 days of aging. Of the treatments evaluated, volatiles of the 5‐day‐old date jaggery solution elicited the highest‐amplitude EAG responses (39%) in A. suspensa females. On the basis of the latter, we tested the attraction response of mature and immature females to date‐jaggery solutions aged over 2 and 4 days in two‐choice flight tunnel bioassays. With both mature and immature females, the 2‐day‐old solution was more attractive than the 4‐day‐old jaggery solution, but significantly more mature females (70% of captures) were attracted to 2‐day‐old jaggery solution. We discuss our results with respect to the improvement of fruit fly lures and attractants by incorporating elements from aged date‐jaggery sugar.     

Effects of adult chill treatments on mate attracting potential for irradiated Mexican fruit fly males

Mexican fruit flies, Anastrepha ludens (Loew), that are reared, sterilized and released for maintenance of a fly free zone in south Texas and for population eradication in California are routinely chilled for 1 to 2 h to facilitate handling and dispersal. While a number of behavioral, physical and physiological characteristics are routinely monitored as part of a quality control program, the effect of the chilling process on the ability of these males to attract wild females has not been evaluated. Tests were performed to compare the mate attracting effectiveness of mass reared, sterilized, chilled males with sterile males that had not been chilled and with recently colonized males. Tests were performed with two recently colonized strains of females in a large cage and males of the three types in plastic cups set on top of the cage. Controls consisted of empty cups. Results showed no significant differences among the three types of males. In tests with the most recently collected (least laboratory adapted) wild strain of females, the chilled males attracted the most females. Fly age and time of day were both significantly related to total attractiveness of males. Number of females responding to males per day increased over the five-day test period. The time period when female response was greatest was variable among the tests.     

Starvation and the Mating Success of Wild Male Mediterranean Fruit Flies (Diptera: Tephritidae)

Recent evidence suggests that the nutritional state of male Mediterranean fruit flies, Ceratitis capitata (Wied.) (medfly), is an important influence on various components of their reproductive biology, including mating success. The objective of the present study was to examine experimentally the effect of temporary starvation on the mating success of wild male C. capitata. Males were maintained on protein–sugar or sugar-only diets, and for each diet we compared the mating success of continuously fed males versus males starved for 18 or 24 h immediately before testing. In trials conducted on field-caged, host trees, males starved for 24 h obtained only about half as many matings as fed males for both diets. However, when the starvation period was 18 h, starved males reared on the protein–sugar diet mated significantly less frequently than fed males, whereas starved males reared on sugar mated as often as fed males. Measurements of male pheromone calling and female attraction revealed that reduced mating success likely reflected the decreased signaling activity of starved males.     

Behavioral evidence for host races in Rhagoletis pomonella flies

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

Behavioral responses of female oriental fruit flies to the odor of papayas at three ripeness stages in a laboratory flight tunnel (Diptera: Tephritidae)

Behavioral responses of adult female oriental fruit flies, Dacus dorsalisHendel, to the odor of papayas from three ripeness classes were studied using a threechoice flight tunnel bioassay. Laboratoryreared flies were allowed to respond freely to any of three papaya odors (mature green, colorbreak to one-fourth ripe, and one-half to full ripe) emanating from identical (spherical) fruit models. Five behaviors were measured in assessing the fly's relative attraction to the odors (number of landings), arrestment (total fly seconds on sphere), fly-fly interactions on the fruit model (maximum and modal fly density), and acceptance for oviposition (total eggs laid). Females showed no significant difference in total fly landings based on all age classes combined. Significant differences were noted among age classes. Females spent more total time on the sphere and showed a higher maximum density and modal fly density to ripe fruit than to green fruit odors. Ovipositional acceptance of fruit models based on the total number of eggs laid in a sphere was greater in response to the ripefruit odor than to the other two odor classes. Olfactorystimulated behavioral responses of females to the odor of ripe papayas were significantly different from the other ripeness classes for all behaviors at 8 days postemergence and then declined in 11-day-old flies. Behavioral responses were greater during the afternoon than in the morning. Observations of wild oriental fruit flies to papayas in the field indicated a preference for residing on riper fruit. The results of this study are discussed with regard to the role of olfactory inputs generated by the odor of ripening fruit on female attraction and oviposition behavior resulting in infestation of papayas by oriental fruit fly.     

Whole body extract of Mediterranean fruit fly males elicits high attraction in virgin females

The search for effective female attractants emanating from the host or body of fruit flies has been an area of intensive research for over three decades. In the present study, bodies of male Mediterranean fruit flies, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), were extracted with diethyl ether or methanol and subjected to gas chromatography–mass spectrometry. Analysis revealed substantial qualitative and quantitative differences between males from a laboratory culture and wild males captured alive in an orchard. Most notably, the hydrocarbon sesquiterpene (±)‐α‐copaene, which is known to be involved in the sexual behaviour of the species, was found in substantial amounts in wild males, but was not detected in laboratory males. In laboratory tests, 15 laboratory or wild male equivalents of diethyl ether extracts or combined diethyl ether and methanol extracts, or, to a lesser extent, methanol extracts alone, were found to attract virgin females. In a citrus orchard, traps baited with combined diethyl ether and methanol extracts of wild males attracted significantly more virgin females than traps baited with various doses of pyranone or blends of other compounds identified in the extracts or reported in the literature, such as ethyl acetate, ethyl‐(E)‐3‐octenoate, and 1‐pyrroline. Traps baited with blends of compounds, however, displayed substantial attractiveness compared to control (non‐baited) traps. These results are important for better understanding the mating system of C. capitata as well as for further improving existing monitoring and control systems.     

Pre-release feeding on yeast hydrolysate enhances sexual competitiveness of sterile male Queensland fruit flies in field cages

Recent laboratory studies of mass‐reared flies in small cages have found that periods of just 24‐ or 48‐h access to yeast hydrolysate can substantially enhance mating performance of mass‐reared male Queensland fruit flies, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) (‘Q‐flies’). Using field cage tests that provide a better approximation of nature, we here investigated whether access to yeast hydrolysate for 48 h after adult emergence improves the ability of male and female mass‐reared, sterile Q‐flies to compete sexually with wild‐type flies that had been provided continuous access to yeast hydrolysate. Mating probability of sterile males was significantly increased by 48‐h access to yeast hydrolysate; sterile males provided 48‐h access to yeast hydrolysate had mating probability similar to that of wild males provided continuous access to yeast hydrolysate, whereas sterile males deprived of access to yeast hydrolysate had much lower mating probability. Unlike males, access to yeast hydrolysate for 48 h did not increase mating probability of sterile female Q‐flies. We instead found that wild females provided continuous access to yeast hydrolysate had higher mating probability than sterile females that did or did not have 48‐h access to yeast hydrolysate. This result raises the possibility that a bisexual Q‐fly strain might operate essentially as a male‐only release when the flies are given access to yeast hydrolysate during a 48‐h pre‐release holding period. Sterile males given access to yeast hydrolysate for 48 h mated significantly earlier in the evening than wild males and, as in other recent studies, this tendency was associated with an increased tendency to mate on the trees rather than the cage walls. There was no evidence of sexual isolation in this study, as wild and sterile mass‐reared flies showed no evidence of preferential mating with their own kind. Further studies are now needed to assess the potential for pre‐release access to yeast hydrolysate to improve sexual performance and longevity of sterile, mass‐reared, Q‐flies in the field.     

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

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

A NO‐CHOICE WIND TUNNEL OLFACTOMETER SYSTEM FOR EVALUATING THE ATTRACTIVENESS OF PLANT VOLATILES TO ADULT HELICOVERPA ARMIGERA (HÜBNER) (LEPIDOPTERA:NOCTUIDAE) *

Abstract A bioassay system to evaluate adult cotton bollworm (CBW), Helicoverpa armigera, behavioral responses to plant volatiles was described. The system was comprised of a no‐choice olfactometer, a wind tunnel, and a wind tunnel room. The olfactometer was designed to provide for the testing and evaluation of volatiles from a variety of natural and synthetic sources. The behavioral responses in this system demonstrated that the volatiles of wilted leaves of Pterocarya stenoptera, Toona sinensis and Populus canadensis, the main materials of which poplar bundles were made, only could attract virgin females but not mated females and males. The volatiles emitted from fresh carrot and celery flowers did not exhibit significant attractiveness to both sexes of CBW. 0.1% phenylacetaldehyde showed significant attraction for virgin females and males. The biological significance of attraction of volatiles from these plants for CBW was discussed.     

Effects of male nutrition on sperm storage and remating behavior in wild and laboratory Anastrepha fraterculus (Diptera: Tephritidae) females

Male physiological condition can affect his ability to modulate female sexual receptivity. Thus, studying this aspect can have biological and practical implications. Here, we examine how male nutritional status affected the amount of sperm stored, remating rate and refractory period of the tephritid fruit fly Anastrepha fraterculus (Wiedemann) females. Both wild and laboratory flies were evaluated. We also examine female sperm storage patterns. Experiments were carried out by manipulating male adult diet and exposing these males to virgin females. Females mated with differently treated males were either dissected to count the amount of sperm stored or exposed to virgin males to determine remating rate and the length of the refractory period. We found that male nutritional status affected the amount of the sperm stored and the renewal of sexual receptivity in wild flies. For laboratory flies, male nutritional status affected the length of the refractory period but not the amount of sperm stored by females. In addition, we report that the ventral receptacle is not an important organ of sperm storage in this species. We conclude that male nutritional condition influences the ability to modulate female sexual receptivity, possibly through a combination of the quantity and quality of the ejaculate. From an applied perspective, providing males with an enriched diet will likely result in increased efficacy of the sterile insect technique.     

Influence of Male Presence on Sexual Maturation in Female Caribbean Fruit Fly, Anastrepha suspensa (Diptera: Tephritidae)

The presence of males was shown to affect the rate of female ovarian development in the Caribbean fruit fly, Anastrepha suspensa (Loew). Virgin females were maintained either in the absence or presence of males. Those sharing a common space with males had either visual contact with the opposite sex or no visual contact. Three strains of Caribbean fruit fly were tested: mass reared strain (flies in colony for more than 20 years); semi-wild strain, flies recently adapted to the laboratory conditions (ca. 12 months); and a wild strain collected in the field. We found that: (1) Mass reared, semi-wild, and wild strains had different female maturation rates, as measured by the presence of mature oocytes, regardless of male presence. (2) Male presence accelerated maturation in wild females and to a lesser extent in semi-wild flies, but had no effect on the long-domesticated mass reared strain. (3) A barrier between female and male cages removed any possibility of visual communication but had no effect on the males’ effect on female maturation. We discuss the adaptive significance of facultative ovarian maturation and the use of male-produced cues to regulate sexual development, and comment on the rapid rate of selection on female maturation under mass-rearing conditions.     

Feeding and foraging of wild and sterile Mediterranean fruit flies (Diptera: Tephritidae) in the presence of spinosad bait

The sterile insect technique (SIT) is used to control wild Mediterranean fruit fly introductions in California and Florida in the U.S. In the past, bait sprays containing malathion proved invaluable in treating new outbreaks or large populations before the use of SIT. Recently, a spinosad protein bait spray, GF-120, has been developed as a possible alternative to malathion, the standard insecticide in protein bait sprays. In this study, protein-deficient and protein-fed Vienna-7 (sterile, mass-reared, "male-only" strain) flies and wild males and females were evaluated to determine the effectiveness of the GF-120 protein bait containing spinosad with respect to bait attraction, feeding, and toxicology. There were no effects of diet or fly type on feeding duration in small laboratory cages. Wild flies, however, registered more feeding events than Vienna-7 males. Flies that fed longer on fresh bait died faster. Protein-deficient flies were more active and found the bait more often than protein-fed flies. Data suggest that adding protein to the diet of SIT flies may decrease their response to baits, therefore, reduce mortality, and thus, allow the concurrent use of SIT and bait sprays in a management or eradication program.