Suppression of melon fly (Diptera: Tephritidae) populations with releases of Fopius arisanus and Psyttalia fletcheri (Hymenoptera: Braconidae) in North Shore Oahu, HI, USA

Field experiments and surveys were conducted to evaluate the efficacy of releasing Fopius arisanus (Sonan) and Psyttalia fletcheri (Silvestri) parasitoids for suppression of Bactrocera cucurbitae (Coquillett) infesting wild Coccinia grandis L. In 2003 and 2004, P. fletcheri releases combined with natural emergence from wild fly populations resulted in better fly suppression, compared to the control site. While P. fletcheri developed freely on melon fly, F. arisanus was less successful at producing its own progeny, yet causing mortality and a twofold decrease in pupae recovered from ivy gourds. Concurrent releases of both parasitoids exerted a compounded suppressive effect on the melon fly population 2–3 times higher than during the pre-release phase. A similar, less obvious, pattern occurred in 2004, due to reduction of the ivy gourd fruit canopy. In 2005, only P. fletcheri was released, with greatly reduced impact, due to ivy gourd destruction and by growers leaving crop culls in fields, producing large numbers of melon flies unaffected by parasitoid releases.     

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.     

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.     

Oviposition response and development of the egg-pupal parasitoid Fopius arisanus on Bactrocera oleae, a tephritid fruit fly pest of olive in the Mediterranean basin

To date, information is wanting with regard to the use of new exotic parasitoids against olive fruit fly, Bactrocera (=Dacus) oleae (Gmelin) (Diptera: Tephritidae), a serious pest of olives Olea europaea L., in the Mediterranean basin. We investigated the oviposition response and developmental biology on B. oleae of Fopius (=Biosteres) arisanus (Sonan) (=Opius oophilus Fullaway) (Hymenoptera: Braconidae), an egg-pupal parasitoid of tephritid fruit flies, never tested before as a potential parasitoid of this host. Our results showed that olive fruits infested with B. oleae eggs exerted a relevant attraction to gravid F. arisanus and represented a stimulus for oviposition. Nevertheless they were not as attractive to female parasitoids as the Mediterranean fruit fly, Ceratitis capitata Wiedemann (Diptera: Tephritidae), eggs infested papaya fruits (Carica papaya L.). In our experimental conditions, F. arisanus completed development in B. oleae within 33 ± 1.7 days (males) and 35 ± 1.6 (females). Increases in host egg to female parasitoid ratios of 1:1, 5:1, 10:1 and 20:1 corresponded with decreases in the percentage of B. oleae parasitisation and host killing but corresponded also with increases in absolute parasitisation. Our findings are discussed in light of possibilities of utilising F. arisanus for biological control of olive fruit fly.     

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。     

Competitor‐free space mediates non‐target impact of an introduced biological control agent

Abstract 1. Enemy‐free space has been shown to mediate host shifts in herbivores, but this has not previously been documented in parasitoids. Also, natural enemies shown to maintain host shifts have always been from higher trophic levels, rather than competitors. 2. In Hawaii, an Australian parasitoid (Diachasmimorpha tryoni) of medflies that loses competition contests to a subsequently introduced Asian parasitoid (Fopius arisanus) has shifted its realised host range to attack non‐target gall flies on lantana. 3. The present study demonstrates experimentally that D. tryoni reproduction is: (i) lower on medflies in coffee when F. arisanus is present than when it is absent; (ii) higher in gall flies on lantana than on medflies in coffee, when F. arisanus is present; and (iii) higher in medflies on coffee than in gall flies on lantana, when F. arisanus is absent. This meets Berdegue et al.’s (Ecological Entomology, 21 , 203–217, 1996) three conditions to confirm the importance of enemy‐free space. 4. In the field, F. arisanus is abundant on medflies, whereas D. tryoni is rare on medflies, but is the dominant parasitoid of lantana gall flies. 5. Competitor‐free space is thus shown to be a key mechanism maintaining an apparent host shift by an introduced biocontrol agent onto a non‐target species.     

The fruit fly parasitoid Fopius arisanus: reproductive attributes of pre-released females and the use of added sugar as a potential food supplement in the field

Fopius (= Biosteres) arisanus (Sonan) (= Opius oophilus Fullaway) (Hymenoptera: Braconidae) is an egg-pupal parasitoid of tephritid fruit flies. Small scale releases of F. arisanus were undertaken in coffee, Coffea arabica L., on the island of Kauai, to evaluate its potential for suppression of fruit fly populations. In conjunction with field releases, assays were conducted to determine the quality of pre-released parasitoids. In addition, the suitability of various sugars as potential food supplements in the field was evaluated. Spermathecal dissection showed that >70% of a total 1.7 million females released in 1998 and 1999 were successfully mated. Moreover, a sperm rating of 2.5–2.6 indicated that the spermathecae of inseminated females were 50% full. There were 85-137 matured eggs (mean =101.7±11.5) in the ovaries of mated or virgin females. When honey was replaced with ripe coffee and water, ripe coffee, or water alone, 10 days after emergence, females remained alive for another 5.5, 5.4, and 4.9 days, respectively. There was corresponding deterioration in the ovaries of gravid females. Mean eggs of honey-deprived females declined from >100 to <70 shortly before their death. Suitability test of various sugars showed that females lived longer when fed with honey (26 days), maple syrup (25 days) or molasses (20 days). Moreover, females provided with these sugars produced mean cumulative progeny per female of 77.2±19, 67±7, and 81±14, respectively, during a 2-week egging period. Our findings provided a basis for development of field release strategy for F. arisanus.     

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.     

Biological Control of Anastrepha spp. (Diptera: Tephritidae) in Mango Orchards through Augmentative Releases of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae)

Diachasmimorpha longicaudata (Ashmead) parasitoids were released by air on a weekly basis over 1600 ha of commercial mango orchards, backyard orchards, and patches of native vegetation, at a density of ca. 940 parasitoids/ha. Releases were made during 2 consecutive years, beginning at flower onset and lasting until the end of the production cycle. Two areas, 7 km apart, were compared. In one area parasitoids were released, whereas the other area was used as a control. During the 2nd year treatments were reversed. Fruit was sampled in commercial mango orchards and in backyard orchards to assess levels of parasitism in fruit fly larvae. Highly significant differences in percentage parasitism were found in release and control zones in backyard orchards. Furthermore, trapping results indicated that D. longicaudata releases were associated with ca. 2.7-fold suppression of Anastrepha spp. populations in backyard orchards. Results suggest that suppression might be affected by environmental conditions and by the parasitoid:fly ratio achieved. Anastrepha obliqua McQuart populations were suppressed more effectively by use of parasitoids than those of Anastrepha ludens Loew, perhaps due to the type of host fruits used by each species. Augmentative parasitoid releases in marginal areas surrounding commercial orchards (backyard orchards, wild vegetation) can substantially suppress fly populations. Through this approach, the number of flies that later move into commercial orchards can be significantly reduced. Such a strategy, when combined with sound orchard management schemes, can allow growers to produce clean fruit without the need to resort to the widespread use of insecticides.     

An Optimized Protocol for Rearing Fopius arisanus, a Parasitoid of Tephritid Fruit Flies

Fopius arisanus (Sonan) is an important parasitoid of Tephritid fruit flies for at least two reasons. First, it is the one of only three opiine parasitoids known to infect the host during the egg stage¹. Second, it has a wide range of potential fruit fly hosts. Perhaps due to its life history, F. arisanus has been a successfully used for biological control of fruit flies in multiple tropical regions^2-4. One impediment to the wide use of F. arisanus for fruit fly control is that it is difficult to establish a stable laboratory colony^5-9. Despite this difficulty, in the 1990s USDA researchers developed a reliable method to maintain laboratory populations of F. arisanus^10-12. There is significant interest in F. arisanus biology^13,14, especially regarding its ability to colonize a wide variety of Tephritid hosts^14-17; interest is especially driven by the alarming spread of Bactrocera fruit fly pests to new continents in the last decade¹⁸. Further research on F. arisanus and additional deployments of this species as a biological control agent will benefit from optimizations and improvements of rearing methods. In this protocol and associated video article we describe an optimized method for rearing F. arisanus based on a previously described approach¹². The method we describe here allows rearing of F. arisanus in a small scale without the use of fruit, using materials available in tropical regions around the world and with relatively low manual labor requirements.     

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).     

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.

     

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.     

Releases of Psyttalia fletcheri (Hymenoptera: Braconidae) and sterile flies to suppress melon fly (Diptera: Tephritidae) in Hawaii

Ivy gourd, Coccinia grandis (L.) Voigt, patches throughout Kailua-Kona, Hawaii Island, HI, were identified as persistent sources of melon fly, Bactrocera cucurbitae (Coquillett). These patches had a low incidence of Psyttalia fletcheri (Silvestri), its major braconid parasitoid natural enemy in Hawaii, and were used to evaluate augmentative releases of P. fletcheri against melon fly. In field cage studies of releases, numbers of melon flies emerging from ivy gourd fruit placed inside treatment cages were reduced up to 21-fold, and numbers of parasitoids were increased 11-fold. In open field releases of P. fletcheri into ivy gourd patches, parasitization rates were increased 4.7 times in release plots compared with those in control plots. However, there was no significant reduction in emergence of melon flies from fruit. In subsequent cage tests with sterile melon flies and P. fletcheri, combinations of sterile flies and P. fletcheri produced the greatest reduction (9-fold) in melon fly emergence from zucchini, Cucurbita pepo L. Reductions obtained with sterile flies alone or in combination with parasitoids were significantly greater than those in the control, whereas those for parasitoids alone were not. Although these results suggest that the effects of sterile flies were greater than those for parasitoids, from a multitactic melon fly management strategy, sterile flies would complement the effects of P. fletcheri. Cost and sustainability of these nonchemical approaches will be examined further in an ongoing areawide pest management program for melon fly in Hawaii.     

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.     

Improving the biological control of leafminers (Diptera: Agromyzidae) using the sterile insect technique

The leafminer Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) is a worldwide pest of ornamental and vegetable crops. The most promising nonchemical approach for controlling Liriomyza leafminers in greenhouses is regular releases of the parasitoid Diglyphus isaea (Walker) (Hymenoptera: Eulophidae). In the current study, we examine the hypothesis that the use of D. isaea for biological control of leafminers in greenhouse crops may be more practical and efficient when supplemented with additional control strategies, such as the sterile insect technique (SIT). In small cages, our SIT experiments suggest that release of sterile L. trifolii males in three sterile-to-fertile male ratios (3:1, 5:1, and 10:1) can significantly reduce the numbers of the pest offspring. In large cage experiments, when both parasitoids and sterile males were released weekly, the combined methods significantly reduced mine production and the adult leafminer population size. Moreover, a synergistic interaction effect between these two methods was found, and a model based on our observed data predicts that because of this effect, only the use of both methods can eradicate the pest population. Our study indicates that an integrated pest management approach that combines the augmentative release of the parasitoid D. isaea together with sterile leafminer males is more efficient than the use of either method alone. In addition, our results validate previous theoretical models and demonstrate synergistic control with releases of parasitoids and sterile insects.     

Effect of host Bactrocera dorsalis sex on yield and quality of the parasitoid Fopius arisanus

This study examines the effect of host Bactrocera dorsalis Hendel (Diptera: Tephritidae) sex on performance of the parasitoid Fopius arisanus Sonan (Hymenoptera: Braconidae) to enable use of a genetic sexing strain (GSS) to transfer this parasitoid to regions where B. dorsalis is not established with lower risk of accidental introduction of the host. Sex ratio and yield of F. arisanus did not significantly differ between the two sexes of a GSS, or between GSS and a standard colony strain. F1 parasitoids of individuals that emerged from the standard colony strain, male GSS hosts and female GSS hosts had similar hatching and parasitization rates on colony eggs. Survivorship of F1 parasitoids with parents from female hosts was significantly greater than for those from male hosts. Using GSS to improve the safety of parasitoid shipments is suggested to be viable, but there may be fitness effects of host sex on F1 parasitoid fitness.     

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.     

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.     

Spatial patterns of sterile Mediterranean fruit fly dispersal

The success of the sterile insect technique (SIT) for the control of the Mediterranean fruit fly or medfly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), depends largely on the ability of sterile flies to spread in the target area and compete with the wild males for wild females. Our objectives in the present study were three‐fold: (1) to evaluate the dispersal ability of sterile male medflies and compare their spatial dispersion patterns with that of wild males, (2) to evaluate how different release methods affect subsequent spatial dispersal, and (3) to determine whether manipulating the pre‐release diet of sterile males affects their dispersal. To achieve these objectives, we conducted three experiments in the field where we quantified and analyzed the spatial and temporal dispersal patterns of sterile medflies and the dispersion of resident wild males. Overall, ca. 5% of the released sterile flies were recaptured 100 m from the release point, and ca. 2% were recaptured 200 m from the release point. The released flies rarely survived longer than 5–7 days. We repeatedly found that the spatial dispersion patterns of sterile males significantly correlated with those of wild males. Release methods strongly affected subsequent fly dispersal in the field as significantly more flies were recaptured following a scattered release vs. a central one. Finally, we show that enriching sterile fly pre‐release diet with protein did not affect subsequent dispersal in the field. We conclude that sterile males are able to match the dispersion patterns of wild males, an outcome that is highly important for SIT success. Large releases from central points distant from each other may leave many areas uncovered. Accordingly, scattered releases, repeated twice a week, will provide better coverage of all available aggregations sites. The spatial performance of protein‐fed males suggests that pre‐release diet amendments may be used without detriment as a sexual stimulant in SIT programs.