Effect of host diet and adult parasitoid diet on egg load dynamics and egg size of braconid parasitoids attacking Anastrepha ludens

The quantity and quality of host nutrients can affect fitness‐related traits in hymenopteran parasitoids, including oogenesis. The present study tested the prediction that a high host quality will influence oogenesis‐related traits positively in synovigenic parasitoids, and that a high‐quality adult parasitoid diet can positively affect the same parameters, potentially compensating for development on low‐quality hosts. Four braconid parasitoid species with contrasting life histories are reared on a low‐quality diet [Anastrepha ludens Loew (Diptera: Tephritidae) larvae reared on mango] or a high‐quality (artificial) diet. Adult parasitoids are provided with a high‐quality (honey ad libitum), moderate‐quality (honey every other day) or low‐quality (guava pulp) diet. Generalist species that encounter high variation in host quality naturally are predicted to be more flexible in dealing with nutrient shortfalls than specialist species. By contrast to the predictions, low‐quality hosts yield parasitoids with higher egg loads in two species: Opius hirtus Fisher and Diachasmimorpha longicaudata Ashmead. However, as predicted, a high‐quality adult diet exerts a positive effect on egg load (Utetes anastrephae Viereck), egg size (Doryctobracon crawfordi Viereck) and egg maturation rate (D. longicaudata, O. hirtus and U. anastrephae). The generalist D. longicaudata varies in egg load and maturation rate depending on host quality and adult diet, respectively. Evidence of the combined effect of both factors on parasitoid fertility is presented for the specialist O. hirtus. The theoretical and practical implications of these findings are discussed.     

Age and host effects on clutch size in the Mexican fruit fly, Anastrepha ludens

We document individual and age-specific variation in reproductive output and clutch size of Anastrepha ludens Loew. (Diptera: Tephritidae). The influence of host size, color, and density on clutch size are also examined. Individual and groups of flies were offered artificial hosts composed of agar spheres wrapped in Parafilm. The gross reproduction rate of individual flies was 1000 eggs/female and 165 clutches/female with a range of 1 to 40 eggs/clutch. Mean clutch sizes for these females ranged from 4.5 to 10.6 eggs/clutch. The number of eggs/clutch laid by females held in groups was highly correlated with host size, ranging from about 4.4 eggs/clutch in 2 cm diameter hosts to 12.7 eggs/clutch in 11 cm hosts. Host color, host density, fly density, and fly age did not affect clutch size. This study suggests that variation among females and host size are the principal determinants of clutch size in A. ludens.

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Résumé L'examen a porté sur l'influence du polymorphisme et des modifications dues à l'âge des mouches sur la fécondité et la taille des pontes de A. ludens Loew (Dipt. Tephritidae). L'étude a concerné aussi l'influence de la dimension, de la couleur et de la densité des hôtes sur la taille des pontes. Des mouches isolées ou par groupes ont reçu des hôtes artificiels formés de sphères d'agar enveloppées dans du parafilm. La fécondité brute de femelles isolées a été de 1000 oeufs/femelle et de 165 pontes/femelle, avec une variation de 1 à 40 oeufs/ponte. La taille moyenne des pontes des différentes femelles isolées s'étalait de 4,5 à 10,6 oeufs/ponte. Le nombre d'oeufs/ponte des femelles groupées était fortement liée à la dimension de l'hôte, s'étalant de 4,4 oeufs/ponte pour des hôtes de 2 cm de diamètre à 12,7 oeufs/ponte pour ceux de 11 cm de diamètre. Ni la couleur et la densité des hôtes, ni la densité et l'âge des mouches n'ont influé sur la taille des pontes. Ces résultats suggèrent que les variabilités du comportement des femelles et de la dimension des hôtes déterminent par priorité la taille des pontes de A. ludens.

     

Superparasitism in mass reared Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae), a parasitoid of fruit flies (Diptera: Tephritidae)

Superparasitism frequency and its effects on the quality of mass-reared Diachasmimorpha longicaudata (Ashmead) parasitoids were investigated under laboratory conditions. Percentage of adult emergence, sex ratio, survival, fecundity and flight ability of adult parasitoids that emerged from Anastrepha ludens (Loew) pupae with different levels of superparasitism were determined. A high prevalence of superparasitism was observed. The number of scars per pupa, produced by insertion of the parasitoid ovipositor, ranged from 1 to 30, with an average (±SD) of 8.3 ± 6.2. Adult parasitoid emergence decreased as the level of superparasitism increased. However, the fraction of females rose with increasing superparasitism and the flight ability was lower in adults emerging from pupae with only one scar, compared with adults emerging from superparasitized hosts. Female longevity and fecundity were not affected by superparasitism. Our results support the hypothesis that superparasitism in D. longicaudata might be adaptive, since adults emerging from hosts with moderate levels of superparasitism showed the highest percentage of emergence and there were no significant differences in the other quality control parameters tested. Our findings are relevant to the mass rearing process, where the ratio of hosts to parasitoids can be optimized as well as the distribution of eggs deposited in host larvae. This contributes to efficient mass rearing methods for augmentative biological control programs.     

Effects of fruit fly host, fruit species, and host egg to female parasitoid ratio on the laboratory rearing of Biosteres arisanus

The host suitability of the oriental fruit fly, Bactrocera dorsalis (Hendel), for development of Biosteres arisanus (Sonan), a braconid parasitoid, was compared with three other fruit fly species, namely, Mediterranean fruit fly, Ceratitis capitata Weidemann, melon fly, Bactrocera cucurbitae Coquilett, and Malaysian fruit fly, Bactrocera latifrons (Hendel). In addition, effects of five different fruit species, namely, Carica papaya L. (solo papaya), Musa sapientum (L.) O. Ktze. (apple banana), Mangifera indica (L.) (Haden mango), Terminalia catappa (L.) (false kamani), and Citrus aurantiifolia (Christman) Swingle (common lime), on the parasitization rate of B. dorsalis and sex ratio of parasitoid progenies were evaluated. Effects of host egg to female B. arisanus ratios on parasitoid progeny yields were likewise determined. The host suitability of fruit flies for development of B. arisanus was ranked as: B. dorsalis>C. capitata=B. latifrons=B. cucurbitae. Based on percent parasitization of B. dorsalis, preference of B. arisanus females for host eggs varied with fruit species, however, preferential oviposition displayed by female parasitoids did not influence sex ratios of subsequent parasitoid progenies. Increases in host egg to female parasitoid ratios of 5:1, 10:1, 20:1, 25:1, and 30:1 corresponded with increases in parasitoid progeny yield reaching a plateau at 20:1.     

Influence of Conspecific Presence, Experience, and Host Quality on Oviposition Behavior and Clutch Size Determination in Anastrepha ludens (Diptera: Tephritidae)

Females of some insect species adjust the number of ovipositions and clutch size adaptively depending on conspecific density and probably experience. In a series of three experiments, we examined the effect of the presence of conspecifics, host quality, and oviposition experience on oviposition behavior and clutch size determination by females of the polyphagous fruit fly Anastrepha ludens (Diptera: Tephritidae). In the first experiment, we determined that grouped (eight females per cage) A. ludens females tended to visit and oviposit in more hosts than did solitary females probably as a result of stimulation by the presence of conspecifics. We also determined that females with previous oviposition experience visited and oviposited in more hosts than inexperienced ones. Importantly, when females were grouped, we observed significantly more landings on unoccupied hosts (i.e., devoid of flies) than on occupied ones (i.e., with at least one fly on it). However, oviposition experience, and not female density, was the most important factor affecting clutch size. Naive females deposited larger egg clutches than experienced ones. In the second experiment, we found that oviposition experience and host quality (i.e., clean fruit or fruit covered with a host marking pheromone [HMP] extract), influenced clutch size and the decision of females to defend or not defend the host. Clutch size and number of fights were greater on clean than on HMP-marked hosts. In the third experiment, we observed that host quality (i.e., size) played a significant role with regard to the number of female fights, host marking behavior, and clutch size. Specifically, females fought and dragged their aculeus longer on small- and medium-sized hosts than on large ones. But this behavior varied according to whether females were kept alone or grouped. Clutch size was greatest in the largest hosts. Considering all the above, we believe that the observed increase in ovipositional activity by grouped A. ludens females can be attributed to competition through mutual interference and not social facilitation as has been reported in other tephritid species.     

Evaluation of entomopathogenic nematodes against the olive fruit fly, Bactrocera oleae (Diptera: Tephritidae)

Infectivity of six entomopathogenic nematode (EPNs) species against Bactrocera oleae was compared. Similar infection levels were observed when third-instar larvae were exposed to infective juveniles (IJs) on a sand-potting soil substrate. When IJs were sprayed over naturally infested fallen olives, many larvae died within treated olives as well as in the soil; Steinernema feltiae caused the highest overall mortality of 67.9%. In addition, three laboratory experiments were conducted to optimize a time period for S. feltiae field application. (1) Abundance of fly larvae inside fallen olives was estimated over the 2006–2007 season with the highest number of susceptible larvae (3 mm and larger) per 100 olives being observed during December, 2006. (2) S. feltiae efficacy against fly larvae dropped to the soil post-IJ-application was determined. B. oleae added to the substrate before and after nematode application were infected at similar levels. (3) Effect of three temperature regimes (min–max: 10–27, 6–18, and 3–12 °C) corresponding to October through December in Davis, California on S. feltiae survival and infectivity was determined. After 8 weeks, the IJs at the 3–12 °C treatment showed the highest survival rate. However, the cold temperature significantly limited S. feltiae infectivity. Our results demonstrate that B. oleae mature larvae are susceptible to EPN infection both in the soil and within infested olives. Being the most effective species, S. feltiae may have the potential to suppress overwintering populations of B. oleae. We suggest that November is the optimal time for S. feltiae field application in Northern California.     

Mating behavior and male mating success in wildAnastrepha Ludens (Diptera: Tephritidae) on a field-caged host tree

Mating behavior and factors affecting mating success of males were studied using wild Anastrepha ludens on a fieldcaged host tree. The most common courtship sequence had five components: (1) male calls from the underside of a leaf, (2) female arrives to the maleoccupied leaf, (3) male orients to female and stops calling, (4) one or both approach to a face-to-face position 1–3 cm apart, and (5) male mounts female after 1–2 s. Courtship behavior was almost identical to that of laboratoryculture flies observed previously under laboratory conditions. Most malefemale encounters occurred at a height of 1–2m, well inside the outer canopy of the tree. Differential mating success by males occurred. No male mated more than once per day, owing possibly to a very short sexual activity period. Factors favoring mating success of males were survival ability and tendency to join male aggregations and to fight other males. Thorax length and age (9–11 days difference) had no effects on male copulatory success. Overall win/loss percentage was not related to mating success because the males that were most successful at mating fought mostly among themselves, driving their win/loss percentage down. However, these successful males (at mating) won most of their fights against less successful males. Results confirmed a lek mating system: males aggregated, called, and defended territories; territories did not contain femalerequired resources; and females exercised mate choice, apparently through selection of sites within leks.     

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.     

Marking of host fruit by maleRhagoletis boycei Cresson flies (Diptera: Tephritidae) and its effect on egg-laying

MaleRhagoletis boycei flies (Cresson) have been shown previously to guard egg-laying punctures on host walnut (Juglans major) fruit. During their tenure,R. boycei males were observed to dip the posterior portion of their abdomen repeatedly toward the fruit surface. Closeup video analysis revealed that, during this dipping behavior, a clear viscous substance emanating from the cercus was deposited directly on the fruit. Laboratory assays indicated that the rate of abdomen-dipping was significantly higher on fruit bearing simulated egg-laying punctures than on control fruit and significantly higher in the area of the puncture than elsewhere on the fruit. In a final experiment, the occurrence of abdomen-dipping in the vicinity of a puncture was manipulated independently on separate punctures on the same fruit. On such fruit and in the absence of males, previously mated, reproductively mature females attempted oviposition significantly more often in or near a puncture exposed to males than in or near an unexposed control puncture. Possible functions of the putative male-marking behavior from the perspectives of both male and female are discussed.     

Enhancing male sexual success in a lekking fly (Anastrepha suspensa Diptera: Tephritidae) through a juvenile hormone analog has no effect on adult mortality

While defending lek-territories, male Anastrepha suspensa (Loew) produce chemical, acoustic and visual courtship signals. In the laboratory and under semi-natural conditions, topical application of the juvenile hormone analog methoprene doubles pheromone production and subsequently doubles sexual success. However, sexual signals and interactions are likely to be physiologically expensive and so result in higher male mortality. Comparison of males kept in isolation for 35 days, but provided daily with a potential mate or a rival male, revealed that both male- and female-interactors shortened focal-male lifespan. In addition, focal males were either treated with methoprene or not, then either provided with protein in their sucrose-based diet or not. Protein proved to similarly double sexual success and also resulted in longer male life spans in all of the interactor-categories. However, there was no evidence that methoprene induced hypersexuality resulted in higher rates of mortality, i.e., the longevity of males treated with methoprene did not significantly differ from untreated males in the same interactor/diet categories. This apparent lack of costs to a putatively sexually selected signal is unexpected but presents an opportunity to increase the sexual competence of sterile flies with few consequences to their survival following mass-release.     

Geissospermum argenteum (Angiosperma: Apocynaceae): a reservoir of parasitoids of fruit flies (Diptera: Tephritidae) in an upland forest in the Brazilian Amazon

1. This work aimed to assess the potential of Geissospermum argenteum Woodson (Apocynaceae) as a reservoir of fruit fly (Diptera: Tephritidae) parasitoids (Hymenoptera) in an upland forest in Brazil.
2. The study was conducted in a remnant of primary upland forest, in which were distributed 10 plots (0.5 ha each). All individuals of G. argenteum with diameters greater than 5 cm at a height of 1.3 m were inventoried. Two trees per plot were selected to assess the total production of fruit. Collectors were installed under the canopy projection of each tree. The fruits were counted and weighed in the field and were transported to the laboratory, to determine the dry mass. Samples of fruits were collected to investigate their infestation by fruit flies and percentage of parasitism.
3. A total of 394 specimens of G. argenteum were identified in the 5 ha investigated area, of which only 126 produced fruits during the 12 months of the assessment. Production ranged from 59 to 2478 fruits per plant (0.59 and 19.73 kg of fresh mass, respectively). Fifty-nine fruit samples were collected from 44 plants, from which 7703 puparia were obtained. Only Anastrepha atrigona Hendel emerged from the puparia. The mean infestation rate was 554.9 puparia/kg of fruit (0.4–1628.6 puparia/kg). Four species of parasitoids were obtained, with mean percentage of parasitism of 2.27% (0.23–9.01%).
4. Our results show that G. argenteum is a reservoir of fruit fly parasitoids, highlighting the importance of its conservation in native habitats, as well as its multiplication in areas close to commercial orchards.

     

Male courtship success and female discrimination in the mediterranean fruit fly (Diptera: Tephritidae)

The mating success of individually marked male Mediterranean fruit flies was monitored over 6 consecutive days in the laboratory. Mating frequency was nonrandom, as the numbers of both males that failed to mate and males that mated many (more than four) times were much higher than expected by chance alone. Differential mating success resulted in part from intermale variation in activity level. Male copulatory success was positively correlated with the numbers of courtships performed, attempted copulations (mountings), and females courted. Male-male aggression, on the other hand, and a negligible effect on male mating success. Female choice also appeared to influence male mating frequency. Fewer than 10% of courtships resulted in mating, and in most cases females terminated courtship by simply moving away from the male. Females that did mate generally selected males having higher copulatory scores than previously rejected males.     

Longevity of multiple species of tephritid (Diptera) fruit fly parasitoids (Hymenoptera: Braconidae: Opiinae) provided exotic and sympatric-fruit based diets

While adult parasitic Hymenoptera in general feed on floral and extrafloral nectars, hemipteran-honeydews and fluids from punctured hosts, Diachasmimorpha longicaudata (Ashmead), an Old World opiine braconid introduced to tropical/subtropical America for the biological control of Anastrepha spp. (Tephritidae), can survive on fruit juices as they seep from injured fruit. An ability to exploit fruit juice would allow such a parasitoid to efficiently forage for hosts and food sources simultaneously. Two New World opiines, Doryctobracon areolatus (Szepligeti) and Utetes anastrephae (Viereck), are also prominent Anastrepha parasitoids and are roughly sympatric. All three species were provided with: (1) pulp and juice diets derived from a highly domesticated Old World fruit (orange, Citrus sinensis L.) that is only recently sympatric with the Mexican flies and parasitoids and so offered little opportunity for the evolution of feeding-adaptations and (2) a less-domesticated New World fruit (guava, Psidium guajava L.), sympatric over evolutionary time with D. areolatus and U. anastrephae. Both sexes of D. longicaudata died when provided guava pulp or juice at a rate similar to a water-only control. D. areolatus and U. anastrephae, presumably adapted to the nutrient/chemical constituents of guava, also died at a similar rate. Survival of all three species on orange pulp and juice was greater than on water, and often equaled that obtained on a honey and water solution. In confirmatory experiments in Mexico, D. areolatus and U. anastrephae, as well as other tephritid parasitoids Doryctobracon crawfordi (Viereck) and Opius hirtus (Fisher), all died at a significantly higher rates when provided guava in comparison to a honey and water diet. Such a result is likely due to guavas being repellent, innutritious or toxic. D. longicaudata clearly consumed guava juice tagged with a colored dye. Dilutions of orange and guava juice resulted in shorter lifespans than dilutions of orange juice and water demonstrating that there while diluted orange juice provided nutrition the addition of guava created toxicity. Given the differences in fruit-food quality, adult opiine food sources would not be obtainable at all oviposition sites and in the case of guava, more additional sites and foraging for food than previously postulated may be required.     

Reproductive behaviour of the South American fruit fly Anastrepha fraterculus (Diptera: Tephritidae): laboratory and field studies

Abstract. The mate choice, courtship and oviposition behaviour of laboratory-reared and field-collected Anastrepha fraterculus (Wied.) were compared. In laboratory cultures in Southampton the duration of male calling activity in small leks increased gradually from 1-2h at 5 days old to up to 7 h at 10 days. This finding correlates with previous reports on the time at which male salivary glands, which are believed to produce sex pheromone, are fully developed. Wild flies which emerged from infested fruits in Brazil began to oviposit on the day they mated, whereas in laboratory flies oviposition began 1 day following the first mating. Both types of fly usually defended their position on a particular fruit throughout the day, and re-mated with either virgin or mated males. There was no significant difference in mating duration. Females did not copulate before the mean age (±SE) of 16.8±0.9 days. For both types of flies mating initiation occurred in the first 2h of photophase, with virgin females choosing mainly mated males. The average number of matings in the laboratory was three for females and four for males, and the interval between matings in females was significantly increased after the second mating. It is suggested that the tendency of virgin females to mate with mated males will lead to increased fitness, as males are on average 48 days old at their second mating. The potential life span of around 200 days for both sexes would allow adults to bridge the gap between seasonally available fruits in warm-temperate and sub-tropical South America.     

Attraction of Mexican Fruit Flies (Diptera: Tephritidae) to Grapefruit: Enhancement by Mechanical Wounding of and Experience with Grapefruit

Wild strain, mated, female Mexican fruit flies, Anastrepha ludens (Loew), with no prior experience with fruit (naive), were not attracted to grapefruit, a preferred cultivated host, in wind tunnel experiments. Naive, mated laboratory strain females were attracted. Prior experience with grapefruit increased attraction of both laboratory and wild strains. More females were attracted to fruit with peel damage than to undamaged fruit, indicating that fruit odor mediated attraction. More naive than experienced females attempted to oviposit on the sides of the wind tunnel. Experienced laboratory males, but not wild males or naive males, were attracted to grapefruit. Attraction and oviposition behavior of laboratory flies was greater than that of wild flies.     

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

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

Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

Pheromone deposition on leaf territories by male Caribbean fruit flies,Anastrepha suspensa (loew) (Diptera: Tephritidae)

Male Caribbean fruit flies apply everted anal membranes associated with pheromone glands to the substrate while on leaf territories. There is a peak in abdominal dipping at the onset of the photoperiod, followed by a decline and then a more extensive period of activity in the second half of the photoperiod. During peak signaling periods, most males had a distinct set of lateral abdominal phermone glands protruded for most of the time. However, the frequency of anal pheromone gland dipping varied considerably over time. Episodes of wing fanning (which may disperse pheromones) and abdomen dipping coincide during a male's tenure on a host leaf. Host-plant leaves were contained for 48 h with signaling males or immature females or kept without insects. Several, though not all, of the constituents of the pheromone were found on leaves within 1 h of removing signaling males. None of these compounds occurred on either type of control leaf. Mature virgin female flies were more likely to be found in contact with host leaves that had been previously exposed for 48 h to the activities of sexually mature males than with unexposed control leaves. There was no difference in the reaction of mature virgin female flies to unexposed control leaves or leaves previously exposed to the activities of other sexually immature female or male flies for 48 h. Presumably, the deposition of pheromones by mature males accounts for the difference in reactions.     

Reproductive biology of Fopius ceratitivorus (Hymenoptera: Braconidae), an egg–larval parasitoid of the Mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae)

The reproductive biology of Fopius ceratitivorus Wharton, a recently discovered African parasitoid, was studied in quarantine in Hawaii to facilitate its mass production for biological control of the Mediterranean fruit fly, Ceratitis capitata. Mean longevity of host-deprived and ovipositing females was 17.3 ± 0.9 d and 16.2 ± 0.5 d, respectively. Ovarian maturation peaked at 61.6 mature eggs per female on the fifth day after eclosion and declined thereafter. Mean number of offspring produced per day by mated females was 5.1 ± 0.4, and realized fecundity expressed as total eggs deposited during the female’s life time was 107.8 ± 12.8. Females were more attracted, to and reproduced significantly more, in fruit substrates containing odors of adult flies and eggs rather than fruit substrates artificially inoculated with fly eggs. Our findings suggest that F. ceratitivorus is a promising new parasitoid for biological control of C. capitata in Hawaii.