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).     

Population dynamics of three Bactrocera spp. fruit flies (Diptera: Tephritidae) and two introduced natural enemies,Fopius arisanus (Sonan) and Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae), after an invasion by Bactrocera dorsalis (Hendel) in Tahiti

Oriental fruit fly, Bactrocera dorsalis (Hendel), invaded French Polynesia in 1996. In 2002 a natural enemy, Fopius arisanus (Sonan), was released and established. By 2009 mean (±SD) F. arisanus parasitism for fruit flies infesting Psidium guajava (common guava), Inocarpus fagifer (Polynesian chestnut) and Terminalia catappa (tropical almond) fruits on Tahiti Island was 64.8 ± 2.0%. A second parasitoid, Diachasmimorpha longicaudata (Ashmead), was released and established in 2008. Although widespread, parasitism rates have not been higher than 10%. From 2003 (parasitoid establishment) to 2009 (present survey) numbers of B. dorsalis, Bactrocera tryoni (Froggatt), Queensland fruit fly, and Bactrocera kirki (Froggatt) emerging (per kg of fruit) declined. For example, for P. guajava there was a decline of 92.3%, 96.8%, and 99.6%, respectively. Analysis of co-infestation patterns (1998–2009) of B. dorsalis, B. tryoni, and B. kirki, suggest B. dorsalis is now the most abundant species in many common host fruits. Establishment of F. arisanus is the most successful example of classical biological control of fruit flies in the Pacific outside of Hawaii and can be introduced if B. dorsalis spreads to other French Polynesian islands, as was the recent case when B. dorsalis spread to the Marquesas Islands. These studies support F. arisanus as a prime biological control candidate for introduction into South America and Africa where Bactrocera carambolae Drew and Hancock and Bactrocera invadens Drew, Tsuruta, and White, respectively, have become established.     

Natural establishment of a parasitoid complex on Bactrocera latifrons (Diptera: Tephritidae) in Hawaii

Bactrocera latifrons (Hendel) (Diptera: Tephritidae) is the most recent of four tephritid fruit fly species accidentally introduced into Hawaii. Although parasitoids have been released against other tephritid fruit fly species and have shown partial success in Hawaii, no parasitoids were released until 2004 to suppress populations of B. latifrons. The present study was conducted to document the parasitoid complex that has naturally established against B. latifrons in Hawaii and to assess whether there is a need for improving the biological control of this species. Based on ripe turkeyberry (Solanum torvum Sw) fruit collections over three consecutive years B. latifrons was the dominant tephritid fruit fly infestating turkeyberry at all four sites surveyed, across three major islands in Hawaii. The overall percentage parasitism of B. latifrons ranged from a low of 0.8% (Hana, Maui) to a high of 8.8% (Kahaluu, Oahu). Five primary parasitoid species were recovered from individually held B. latifrons puparia: Fopius arisanus (Sonan), Psyttalia incisi (Silvestri), Diachasmimorpha longicaudata (Ashmead), D. tryoni (Cameron), and Tetrastichus giffardianus Silvestri. F. arisanus was the predominant parasitoid at three of the four sites. Low levels of parasitism suggest that there is a need to improve biological control of B. latifrons, to minimize chances of this species causing economic impacts on crop production in Hawaii. We discuss the possibility of improving biological control of B. latifrons through augmentative releases of F. arisanus or introduction and release of specific and efficient new parasitoid species.     

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.     

Host species and vegetable fruit suitability and preference by the parasitoid wasp Fopius arisanus

Parasitoids that oviposit in a concealed host inside a plant part need to be able to find both the plant and the host. Egg parasitoids of fruit‐infesting Tephritidae need to assess the oviposition site based both on the host egg and the infested fruit. Infestation by Tephritidae fruit flies threatens fruit and vegetable production. Management methods have been implemented including biological control, using Fopius arisanus Sonan (Hymenoptera: Braconidae). The parasitism by F. arisanus in three Tephritidae flies in vegetable fruits was investigated. Laboratory assays were conducted to assess the parasitoid's preference and survival. Zucchini, sweet pepper, and tomato were artificially infested with eggs of Bactrocera dorsalis Hendel, Ceratitis capitata Wiedemann, and Ceratitis cosyra Walker (all Diptera: Tephritidae), then exposed to mated naïve F. arisanus females in a 20:1 egg:parasitoid ratio. Parasitoid behavioral activities (resting, antennating, probing, ovipositing) were observed on the infested fruits. Parasitism rate was determined by dissection of fruit fly eggs under a stereomicroscope. Behavioral activities of F. arisanus differed between all the fruits when infested with B. dorsalis or C. cosyra eggs but differed only between some of the fruits when infested with C. capitata. Fopius arisanus preferred B. dorsalis over C. capitata and C. cosyra, with a parasitism rate 2× higher on B. dorsalis compared to the Ceratitis species. Preference for fruits was dependent on the infesting fruit fly. The emergence of F. arisanus was higher with B. dorsalis than with Ceratitis spp. Although B. dorsalis completed its development earlier than Ceratitis spp., host fly species did not affect the developmental time of F. arisanus. We discuss the significance of F. arisanus preference in relation to naturally occurring Tephritidae infestations. We also discuss whether some fruits might constitute a refuge for Tephritidae flies and whether this will affect the current biological control efforts against B. dorsalis.     

Biological attributes of three introduced parasitoids as natural enemies of fruit flies,genus Anastrepha (Diptera: Tephritidae)

The biological attributes of three introduced species of parasitoids which attack the fruit fly Anastrepha ludens were evaluated. Larvae and eggs of A. ludens were exposed to larval parasitoids Diachasmimorpha longicaudata and D. tryoni and the egg parasitoid Fopius arisanus. Parasitoid longevity and fecundity were determined using larvae and eggs of A. ludens. Likewise, the parasitism rates of these parasitoid species in infested host fruits were recorded. The intrinsic rate of increase for F. arisanus was 0.1019 followed by D. tryoni with a rate of 0.1641 and D. longicaudata with the highest rate of 0.2233. Although F. arisanus females had the highest levels of fecundity, only 50% of them remained alive until reproductive age. These results in combination with the longer generation time (in comparison with D. longicaudata and D. tryoni), can be considered as the most important factors explaining F. arisanus reduced rate of increase. However, we note that oviposition activity caused egg mortality which reduced Anastrepha egg hatch by ca. 20%. This result suggests that F. arisanus has a high potential as a natural enemy of A. ludens, in accordance with our research efforts to develop a new F. arisanus strain specialized for development in Anastrepha eggs. The results show that D. tryoni is not a good candidate for biological control of Anastrepha. Meanwhile, D. longicaudata continues to be the most important exotic parasitoid for suppression of Anastrepha fruit fly populations.     

Prevalence and evolutionary history of endosymbiont Wolbachia (Rickettsiales: Anaplasmataceae) in parasitoids (Hymenoptera: Braconidae) associated with Bactrocera fruit flies (Diptera: Tephritidae) infesting carambola

Wolbachia endosymbiont is a maternally inherited bacteria that infects a wide range of hosts, including parasitoids and their respective hosts. In this study, a total of 171 individuals of braconid endoparasitoids, consisting of Fopius arisanus, F. vandenboschi, Diachasmimorpha longicaudata, Psyttalia sp.1, Psyttalia sp.2, P. fletcheri and P. incisi, and their host tephritid fruit flies of Bactrocera dorsalis, B. papayae and B. carambolae infesting carambola were screened molecularly by the Wolbachia surface protein (wsp) gene. Interestingly, 21 (24.14%) wsp gene sequences were successfully isolated from 87 braconid samples tested, showing a low infection rate of Wolbachia. However, despite the close ecological contact between parasitoids and their hosts, none of the tephritid individuals were infected by Wolbachia. A comparison of wsp and host mitochondrial cytochrome c oxidase subunit I (COI) sequences found that braconids did not cluster in connection with Wolbachia infection, suggesting that selective sweep has not yet occurred because Wolbachia may have recently infected the braconid populations in Peninsular Malaysia (≈0.1 MYA). Despite of relatively recent infections of Wolbachia, the history of Wolbachia infection into F. arisanus populations of Peninsular Malaysia is complex, involving at least two independent occasions of infection and two secondary losses.     

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.     

Opiine parasitoids (Hymenoptera: Braconidae) and biological control of fruit flies (Diptera: Tephritidae) in Australia: Past,present and future

Opiine braconids are parasitoids of the immature stages of frugivorous tephritids. The female wasp lays her eggs into the eggs or larvae of the fruit fly host, where the immature wasp develops before emerging as a next-generation adult from the now dead host pupal case. In support of a new generation of Australian fruit fly parasitoid research, this paper comprehensively reviews what is known about the Australian fruit fly infesting opiines. Based on the most recent taxonomic revision 11 fruit fly infesting opiine species are documented to occur in Australia, but we consider as doubtful the record for Diachasmimorpha longicaudata and consider the record for Fopius illusorius to be tentative without further collections. We identify that the systematics and taxonomy of the Australian native fruit fly infesting opiines are in urgent need of further work. The history of fruit fly biological control in Australia is comprehensively reviewed, including the export of native Australian opiines for fruit fly control elsewhere in the world. Australia was actively involved in three major classical biological control programmes against fruit flies from the turn of the 1900s until the mid-1960s. Despite the introduction of 11 opiine species, plus numerous other natural enemies, only Fopius arisanus established in eastern Australia, where in South-east Queensland it can now cause between 30 and 40% mean parasitism. In addition to the exotic F. arisanus, the native species Diachasmimorpha kraussii and Diachasmimorpha tryoni also cause fruit fly parasitism in agriculturally important crops: both species have also been liberated widely outside of Australia for fruit fly control. Other Australian opiines have not been reared from flies infesting commercial crops and appear biologically restricted to the fruits and environs of Australian east-coast rainforests. The biology literature for D. tryoni and D. kraussii is comprehensively reviewed, while for F. arisanus, already reviewed elsewhere, key literature only is covered. Forward looking, we consider the potential for inoculative or inundative releases of opiines in areas where they do not currently occur to be good, while conservation biological control may help to increase the impacts of parasitoids in areas where they are already established.     

Implications of host mortality on the economics of Fopius arisanus (Hymenoptera: Braconidae) mass rearing

Fopius (= Biosteres) arisanus (Sonan) (= Opius oophilus Fullaway) is an egg-pupal parasitoid of tephritid fruit flies. Our breakthrough in the development of a laboratory-adapted strain of F. arisanus facilitated insectary rearing of parasitoids in large numbers. First colonized in captivity in 1989, F. arisanus has been reared routinely on the oriental fruit fly, Bactrocera (= Dacus) dorsalis (Hendel), its natural host. Parasitization by F. arisanus results in the latent death of developing hosts. Host mortality data are presented in a life table to estimate cost of parasitization on the economics of F. arisanus mass rearing. Percent kills of 10, 32, and 76% exerted by F. arisanus on the egg, larval, and pupal stages of the rearing hosts, respectively, resulted in a mean parasitoid recovery of 23.6%. The production cost was estimated by integrating life table data with the costs of rearing supplies and materials and personnel-hours requirements. Production of 1 million parasitoid adults requires the exposure of 4.2 million B. dorsalis eggs (= propagation hosts). We estimated the cost of producing 1 million parasitoids to be $2,363.30 and identified the most expensive aspects of F. arisanus rearing. Recommendations for streamlining the rearing process to reduce costs are provided.     

Effect of different dietary resources on longevity, carbohydrate metabolism, and ovarian dynamics in two fruit fly parasitoids

A growing body of literature reports the importance of non-prey food sources in boosting fitness of arthropod natural enemies, thus further contributing to their pest control efficacy. Although resources such as nectar, pollen, or honeydew have received a fair amount of research attention, little is known about the role of fruit juices. Under natural conditions, Tephritid fruit fly parasitoids enjoy ample access to fallen or damaged fruits and their saccharide-rich juices, and wasp fitness can potentially benefit in multiple ways from access to these resources. In this study, we compared the effect of fruit juice with other food resources on multiple fitness parameters in parasitoids that commonly forage on fallen, damaged fruits: the braconid Diachasmimorpha longicaudata and figitid Aganaspis pelleranoi. Parasitoids were subject to simple or combined diets of guava juice (Psidium guajava), honey and pollen, and their effect on wasp longevity, ovarian dynamics and (body) carbohydrate levels was assessed. For both species, adult longevity proved lowest on simple diets of water, guava juice, or pollen, while greatest longevity was attained on honey or combined diets. For D. longicaudata, egg load did not differ between the various diets, while A. pelleranoi egg load was higher for individuals that had access to honey or pollen, but did not differ between newly emerged wasps and those fed guava juice. In both parasitoid species, total sugars, fructose, and glycogen levels were highest in wasps fed with honey or combined diets and lowest under (simple) guava juice, pollen, or water diets. In conclusion, D. longicaudata and A. pelleranoi attained superior longevity and body nutrient levels with access to high-sucrose sugar sources, such as honey, but benefited comparatively little from access to guava juice. Our work hints the role of high-sucrose foods such as (extra-) floral nectar or artifical sugar sprays in boosting fitness of fruit fly parasitoids. We further discuss the relevance of these findings for fruit fly biological control, in crops such as guava.     

Effect of Two Opiine Parasitoids (Hymenoptera: Braconidae) Introduced for Fruit Fly Control on a Native Hawaiian Tephritid,Trupanea dubautiae(Diptera: Tephritidae)

Diachasmimorpha longicaudata(Ashmead) andPsyttalia fletcheri(Silvestri) are opiine parasitoids introduced into Hawaii for control of the Oriental fruit fly,Bactrocera dorsalis(Hendel) and the melon fly,Bactrocera cucurbitae(Coquillett), respectively. Both species have recently been mass-reared and released for research in augmentative biocontrol programs. Laboratory and field sleeve cage experiments were conducted to investigate the potential impact of mass-producedD. longicaudataandP. fletcherion a native Hawaiian tephritid,Trupanea dubautiae(Bryan), infesting the flowerheads of the native composite shrubDubautia raillardioidesHillebrand. Gravid females ofD. longicaudataandP. fletcheriwere confined with bloomingD. raillardioidesflowerheads infested with late instarT. dubautiae.BothD. longicaudataandP. fletcherilacked positive oviposition responses toT. dubautiaelarvae in infested flowerheads and caused neither parasitism nor mortality to the flies. However, when larvae were removed from the flowerheads and presented in screened dishes containing artificial diet of the parasitoids' normal rearing hosts (B. dorsalisandB. cucurbitae), bothD. longicaudataandP. fletcherireadily oviposited in the test larvae. Oviposition byD. longicaudatadid not significantly affect the percentage pupation ofT. dubautiae,but did reduce the emergence of adult flies. Oviposition byP. fletcherisignificantly reduced both pupation and adult fly emergence. All progeny of both parasitoid species died as eggs or first-instar larvae. Results from our experiments demonstrate that biological control programs targeted against frugivorous tephritid pests byD. longicaudataandP. fletcherihave no harmful impact on flowerhead-infestingT. dubautiae.     

Competition between Diachasmimorpha kraussii and Fopius arisanus in Bactrocera tryoni: does native parasitoid-host association matter?

In Australia Fopius arisanus (Sonan) is an established, but exotic fruit fly egg-larval-pupal parasitoid which co-occurs with Diachasmimorpha kraussii (Fullaway), a native larval-pupal fruit fly parasitoid: both attack the native fruit fly Bactrocera tryoni (Froggatt). In interactions involving evolutionary novel host-parasitoid associations, F. arisanus consistently out-competes other parasitoid species, including D. kraussii. However, in fruit fly-parasitoid systems where there is co-evolutionary history between parasitoids and their hosts, competitive hierarchies can vary. In this study we investigated the outcome of competition between F. arisanus and D. kraussii within B. tryoni, to test whether the close evolutionary relationship between D. kraussii and B. tryoni might circumvent the competitive advantage of F. arisanus. Consistent with previous research, and despite the evolutionary relationship, dissection of multiparasitized B. tryoni larvae showed that D. kraussii was invariably suppressed by F. arisanus. A total of 47% and 74% of the eggs of D. kraussii in presence of F. arisanus were killed within a span of 24 h and 48–72 h, respectively. However, parasitoid emergence from fruit fly hosts exposed sequentially to F. arsianus and D. kraussii suggest that D. kraussii females are able to discriminate hosts already parasitized by F. arisanus. Results show that the co-evolutionary relationship between B. tryoni and D. kraussii does not help overcome the early-acting advantage of the egg parasitoid F. arisanus. Though F. arisanus may not have completely displaced D. kraussii in its native habitat, simultaneous inundative releases of these two parasitoid species (currently under consideration) might not help increase B. tryoni parasitism levels.

     

应用寄生蜂和不育雄虫防控田间橘小实蝇

食品安全已成为当今社会广受关注的问题。尽管国内的一些学者就阿里山潜蝇茧蜂[Fopius arisanus(Sonan)]和橘小实蝇SIT技术分别开展了不同程度的研究,然而关于应用该蜂和不育雄虫对橘小实蝇[Bactrocera dorsalis(Hendel)]进行田间联合防治的研究还未见相关报道。为此,为探讨天敌昆虫和雄性不育技术对橘小实蝇的田间综合治理(IPM)技术,开展了阿里山潜蝇茧蜂和橘小实蝇不育雄虫的田间释放试验:首先应用性引诱剂降低田间野生雄虫数量;而后,释放橘小实蝇不育雄虫,进一步降低田间橘小实蝇种群数量;最后,释放阿里山潜蝇茧蜂对田间残余的橘小实蝇卵进行追踪寄生,以达到持续控制的目的。通过监测整个防控期田间橘小实蝇的种群数量以及果实受害率和防治效果作为评价指标评价该防治方法的综合防治效果。结果表明:应用阿里山潜蝇茧蜂、橘小实蝇不育雄虫并配合引诱剂的综合措施对橘小实蝇的种群数量有明显控制作用,田间总体防治效果可达90%左右。此外,提出完善橘小实蝇田间防控措施的建议,并提出该研究的不足以及有待进一步开展研究的方向。     

The effect of four fruit species on the parasitization rate of Anastrepha fraterculus (Diptera: Tephritidae,Trypetinae) by Diachasmimorpha longicaudata (Hymenoptera: Braconidae,Opiinae) under laboratory rearing conditions

In the laboratory, the effect of host fruit species Citrus paradisi, C. aurantium, Prunus persica, and Psidium guajava on A. fraterculus parasitism by Diachasmimorpha longicaudata was studied. The number of ovipositor-probing events and the probing-time were documented to evaluate the role of fruit chemistry, and epicarp and mesocarp thickness, respectively. The relationship between the parasitization rate and fruit size in particular plant species was analyzed by applying a simple regression. Results showed that guava and peach yielded significantly more parasitoids than both Citrus spp. Probably, the parasitization rate of D. longicaudata on A. fraterculus would be influenced in part by chemical and physical factors from fruit species.     

Intra- and interspecific competition by Fopius arisanus and Diachasmimorpha tryoni (Hymenoptera: Braconidae), parasitoids of tephritid fruit flies

Fopius arisanus (Sonan) and Diachasmimorpha tryoni (Cameron) are two important solitary endoparasitoids of tephritid fruit flies. The former species attacks host eggs while the latter attacks host larvae, and both species emerge as adults from the host puparium. This study investigated intrinsic competition between these two parasitoids, as well as aspects of intraspecific competition within each species in the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). Parasitization by F. arisanus resulted in direct mortality of host eggs and prolonged development of host eggs and larvae. Superparasitism by F. arisanus was uncommon when mean parasitism per host patch was <50%, but increased with rising rates of parasitism. Superparasitism by D. tryoni was more common. In superparasitized hosts, supernumerary individuals of F. arisanus were killed through physiological suppression, while supernumerary larvae of D. tryoni were killed mainly through physical attack. In multiparasitized hosts, dissections showed that 81.6% of D. tryoni eggs in the presence of F. arisanus larvae died within 3 days, indicating physiological inhibition of egg hatch. Rearing results further showed that F. arisanus won almost all competitions against D. tryoni. The ratio of D. tryoni stings to ovipositions was lower in hosts not previously parasitized by F. arisanus than in parasitized hosts, suggesting that D. tryoni can discriminate against parasitized hosts. The mechanism that F. arisanus employs to eliminate D. tryoni is similar to that it uses against all other larval fruit fly parasitoids so far reported. The results are discussed in relation to the competitive superiority of early acting species in fruit fly parasitoids, and to a possible competitive-mediated mechanism underlying host shift by D. tryoni to attack non-target flies following the successful introduction of F. arisanus in Hawaii.     

Olfactory response of three parasitoid species (Hymenoptera: Braconidae) to volatiles of guavas infested or not with fruit fly larvae (Diptera: Tephritidae)

The olfactory responses of the native parasitoids Doryctobracon areolatus (Szépligeti) and Asobara anastrephae (Muesebeck) and of the exotic parasitoid Diachasmimorpha longicaudata (Ashmead) to guava (Psidium guajava L.) infested or not with fruit fly larvae were evaluated. D. areolatus and D. longicaudata females responded to the odors of uninfested rotting guavas, although D. areolatus was also attracted to fruits at the initial maturation (turning) stage. The females of these species recognized the volatiles of guavas containing Ceratitis capitata (Wied.) larvae. However, in bioassays involving fruits with larvae of different instars, D. longicaudata females were not able to separate between fruits containing C. capitata larvae at the initial instars and larvae at the third instar. In the evaluations of volatiles released by guavas containing C. capitata and Anastrepha fraterculus (Wied.) larvae, the D. longicaudata females were oriented toward the volatiles of fruits containing both host species, but differed significantly from volatiles of guavas containing C. capitata larvae. The D. areolatus females also showed responses to both species, although with a preference for volatiles of fruits containing A. fraterculus larvae. The A. anastrephae females were oriented toward the odors of fruits infested with both fruit fly species. In the shade house, D. longicaudata females were oriented to volatiles of rotting fruits containing larvae or not, but could not significantly differentiate between hosts. D. areolatus females were not attracted toward fruits on the ground in the shade house, regardless of host, suggesting that this parasitoid does not forage on fallen fruits.     

Effect of <Emphasis Type="Italic">Fopius arisanus</Emphasis> oviposition experience on parasitization of <Emphasis Type="Italic">Bactrocera dorsalis</Emphasis>

Fopius arisanus (Sonan) (Hymenoptera: Braconidae) is an important fruit fly parasitoid, successfully introduced in programs of classical biological control around the world. One aspect of its reproductive biology that has received increasing attention is the role of learning on parasitization by individual females. In this study, we examined the effect of previous oviposition opportunities on the reproductive success and progeny sex ratio of F. arisanus on eggs of Bactrocera dorsalis Hendel (Diptera: Tephritidae). Our results indicate that the proportion of eggs parasitized and parasitoid yield both increase with increasing experience, as acquired by individual females via previous oviposition opportunities. These effects are shown to be unrelated to parasitoid age out to three weeks, which we found not to have an effect on parasitization, yield, or sex ratio. We discuss the implications of our results on mass rearing and also in terms of the efficiency of this biological control agent in the field.     

Intrinsic inter-specific competition in a guild of tephritid fruit fly parasitoids: Effect of co-evolutionary history on competitive superiority

Tephritid fruit fly parasitoid guilds are dominated by solitary koinobiont species that attack different host stages, but most emerge as adults from host puparia. Previous studies suggest intrinsic competitive superiority by the egg-attacking parasitoid Fopius arisanus (Sonan) against all larval-attacking parasitoids in Hawaii. In this study, we tested the early-acting competitive superiority prediction in relation to the co-evolutionary history of competition between an egg–larval parasitoid (Fopius ceratitivorus Wharton), and each of three larval parasitoids [Psyttalia concolor (Szépligeti), Diachasmimorpha kraussii (Fullaway), and Diachasmimorpha longicaudata (Ashmead)]. F. ceratitivorus and P. concolor share a common origin (eastern Africa), while D. kraussii is an Australian species, and D. longicaudata is from Southeast Asia. The outcomes of intrinsic competition between the egg-attacking parasitoid and each of the three larval-attacking parasitoids within their common host, the Mediterranean fruit fly Ceratitis capitata (Wiedemann) were compared. F. ceratitivorus invariably eliminated the co-evolved P. concolor through physiological suppression of the later-attacking parasitoid’s egg development, providing evidence that supports the early-acting-superiority hypothesis. However, F. ceratitivorus was unable to suppress development of the two non co-evolved larval parasitoids. Instead, the larvae of both later-acting parasitoid species physically killed F. ceratitivorus larvae inside the host. The results suggest that co-evolutionary history influences competitive superiority. The evolution of inter-specific competition and its implications for biological control are discussed.     

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.