Newly imported larval parasitoids pose minimal competitive risk to extant egg-larval parasitoid of tephritid fruit flies in Hawaii

Competitive displacement of fruit fly parasitoids has been a serious issue in the history of fruit fly biological control in Hawaii. This concern regarding competitive risk of new parasitoids has led to an overall tightening of regulations against the use of classical biological control to manage fruit flies. Fopius arisanus (Sonan), an egg-larval parasitoid, is the most effective natural enemy of tephritid fruit flies in Hawaii. This study evaluated the competitive risk of two recently introduced larval parasitoids, Diachasmimorpha kraussii Fullaway and Psyttalia concolor (Szépligeti), to F. arisanus attacking the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). Fopius arisanus won almost all intrinsic competitions against both larval parasitoids through physiological suppression of egg development. 83.3% of D. kraussii eggs and 80.2% of P. concolor eggs were killed within three days in the presence of F. arisanus larvae within the bodies of multi-parasitized hosts. The mechanism that F. arisanusemploys to eliminate both larval parasitoids is similar to that it uses against three other early established larval fruit fly parasitoids: F. vandenboschi (Fullaway), D. longicaudata (Ashmead) and D. tryoni (Cameron). It suggests that introduction of these larval parasitoids poses minimal competitive risk to F. arisanus in Hawaii.     

Host preference and host suitability in an egg-pupal fruit fly parasitoid, Fopius arisanus (Sonan) (Hym., Braconidae)

The relative oviposition rate of the parasitoid Fopius arisanus (Sonan) was investigated across three frugivorous tephritid species, Bactrocera tryoni Froggart, Bactrocera jarvisi (Tryon) and Bactrocera cucumis French. Choice and no-choice tests were both used. The suitability of these three species for sustaining larval development and survival to the adult stage was also assessed. Fopius arisanus parasitized all three tephritid species, regardless of the method of exposure, but showed stronger preference for B. tryoni and B. jarvisi over B. cucumis. Superparasitism was extremely rare. Successful development of F. arisanus varied across host species. Bactrocera tryoni yielded significantly more parasitoids than B. jarvisi, but no wasps emerged from B. cucumis puparia. Tests were set up in replicated trials, but results were not homogeneous across trials. We discuss the host relationships of F. arisanus with reference to this variation and in relation to host suitability for larval development.     

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.     

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.     

Potential for areawide integrated management of Mediterranean fruit fly (Diptera: Tephritidae) with a braconid parasitoid and a novel bait spray

The braconid wasp, Fopius arisanus (Sonan), a biological control agent for Mediterranean fruit fly, Ceratitis capitata (Wiedemann), was studied in coffee, Coffea arabica L. Fopius arisanus, comprised 79.3% of the total parasitoids (7,014) recovered from fruits collected at three small coffee farms. Data from seasonal host/parasitoid studies at a large coffee plantation also suggested that the most effective natural enemy of C. capitata in coffee may now reside in Hawaii. The original parasitoids introduced into Hawaii for C. capitata control (Diachasmimorpha tryoni (Cameron), Tetrastichus giffardianus Silvestri, and Dirhinus giffardii Silvestri) are now rare. Abundance of F. arisanus with respect to other parasitoids collected was influenced by elevation (274, 457, 610 m). Fopius arisanus was the dominant parasitoid at all three elevations, Diachasmimorpha longicaudata (Ashmead) occurred consistently, and T. giffardianus was abundant only at low elevation. The impacts on C. capitata and F. arisanus populations of bait sprays containing malathion, spinosad, or phloxine B applied to coffee were also evaluated. All three bait sprays suppressed C. capitata populations. Spinosad and phloxine B bait sprays appeared less harmful to the wasp than malathion. Fopius arisanus offers the potential for areawide management of C. capitata that includes biological control and integration with more environmentally safe chemical controls such as spinosad and phloxine B bait sprays.     

The Effects of Chilling on the Fecundity and Life Span of Mass-reared Parasitoids (Hymenoptera: Braconidae) of the Mediterranean Fruit Fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae)

Suppression of Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann), populations may be achieved through the mass-rearing and augmentative aerial release of opiine braconid parasitoids. Typically, aerial release techniques require up to one hour of chilling of adult parasitoids at temperatures as low as 3.5°C prior to their dissemination. Such chilling potentially could affect the subsequent performance of the insects. Among three species of the genus Diachasmimorpha longicaudata (Ashmead), tryoni (Cameron), and krausii (Fullaway) there was little or no affect of chilling in the laboratory on female longevity, production of daughters, or offspring sex ratio. This is consistent with previous experiments that found chilling to have no discernable effect on the short-term mortality of D. tryoni or on its ability to take flight immediately after aerial release. While there was little effect of chilling on longevity and fecundity in a species from another opiine genus, Fopius arisanus (Sonan), exposure to low temperatures did result in a significantly more male-biased offspring sex ratio.     

Competitive Superiority of Early Acting Species: A Case Study of Opiine Fruit Fly Parasitoids

Koinobiont parasitoids that attack an early host stage may have an advantage in suppressing competing parasitoids that attack later stages of the same host. We examined the competitive interaction between the two most important parasitoids of tephritid fruit flies in Hawaii, Fopius arisanus (Sonan), and Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae). The former species attacks host eggs while the latter attacks host larvae, and both species emerge as adults from the host puparia. F. arisanus physiologically suppressed egg development of D. longicaudata. Over 90% of D. longicaudata eggs died in the presence of F. arisanus larvae within host larvae of either the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) or the oriental fruit fly, Bactrocera dorsalis (Hendel). D. longicaudata appeared not to discriminate against hosts previously parasitized by F. arisanus. The mechanism that F. arisanus uses to eliminate D. longicaudata is similar to that which it employs to eliminate five other larval fruit fly parasitoids so far reported in Hawaii. This suggests that there is a broad competitive superiority of the early acting species in fruit fly parasitoids. We discuss the implication of this in relation to future biological control introductions against tephritid fruit flies.     

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.     

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.     

Toxicity of spinosad in protein bait to three economically important tephritid fruit fly species (Diptera: Tephritidae) and their parasitoids (Hymenoptera: Braconidae)

The feeding toxicity of the natural insecticide spinosad in Provesta protein bait was evaluated for three economically important fruit fly species, the Mediterranean fruit fly, Ceratitis capitata (Wiedemann); the melon fly, Bactrocera cucurbitae Coquillett; and the oriental fruit fly, Bactrocera dorsalis Hendel. Both females and males were evaluated. Spinosad was remarkably similar in toxicity to all three fruit fly species. Male C. capitata (24 h LC50 values and 95% fiducial limits = 2.8 [2.60-3.0] mg/liter spinosad) were significantly, although only slightly more susceptible to spinosadthan females (4.2 [3.8-4.6] mg/liter). Male (5.5 [4.7-6.6] mg/liter) andfemale (4.3 [3.7-4.9] mg/liter) B. cucurbitae were equally susceptible to spinosad. Female (3.3 [3.1-3.6] mg/liter) and male (3.1 [2.9-3.3] mg/liter) B. dorsalis also were equally susceptible to spinosad. Provesta bait containing spinosad also was evaluated against two parasitoids of tephritid fruit flies, Fopius arisanus (Sonan) and Pysttalia fletcheri (Silvestri). These parasitoids did not feed on the bait, so a contact toxicity test was conducted. Significant amounts of mortality were found only after exposure of parasitoids to spinosad-coated glass vials with concentrations > or =500 mg/liter spinosad. Parasitoids were less susceptible than fruit flies to such a degree that use of spinosad in bait spray should be compatible with these parasitoid species. Because the fruit flies tested in this study were so susceptible to spinosad, this product seems to be promising as a bait spray additive and a replacement for malathion for control of these species.     

Opiine parasitoids (Hymenoptera: Braconidae) of tropical fruit flies (Diptera: Tephritidae) of the Australian and South Pacific region

Opiine wasps are parasitoids of dacine fruit flies, the primary horticultural pests of Australia and the South Pacific. A taxonomic synopsis and distribution and host records (44% of which are new) for each of the 15 species of dacine-parasitizing opiine braconids found in the South Pacific is presented. Species dealt with are Diachasmimorpha hageni (Fullaway), D. kraussii (Fullaway), D. longicaudata (Ashmead), D. tryoni (Cameron), Fopius arisanus (Sonan), F. deeralensis (Fullaway), F. ferrari Carmichael & Wharton sp. n., F. illusorius (Fischer) comb. n., F. schlingeri Wharton, Opius froggatti Fullaway, Psyttalia fijiensis (Fullaway), P. muesebecki (Fischer), P. novaguineensis (Szépligeti) and Utetes perkinsi (Fullaway). A potentially undescribed species, which may be a colour morph of F. vandenboschi (Fullaway), is diagnosed but not formally described. Fopius vandenboschi sensu stricto, Diachasmimorpha fullawayi Silvestri, Psyttalia concolor Szépligeti and P. incisi Silvestri have been liberated into the region but are not considered to have established: a brief diagnosis of each is included. Biosteres illusorius Fischer is formally transferred to the genus Fopius. A single opiine specimen reared from a species of Bactrocera (Bulladacus) appears to be Utetes albimanus (Szépligeti), but damage to this specimen and to the holotype (the only previously known specimen) means that this species remains unconfirmed as a fruit fly parasite: a diagnosis of U. cf. albimanus is provided. Psyttalia novaguineensis could not be adequately separated from P. fijiensis using previously published characterizations and further work to resolve this complex is recommended. A key is provided to all taxa.     

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).     

Parasitons of non-target tephritid flies in hawaii: Implications for biological control of fruit fly pests

We surveyed the parasitoid complex associated with four non-pest tephritid flies in Hawaii,Procecidochares alani Steyskal (on the Island of Hawaii), andEutreta xanthochaeta Aldrich,Phaeogramma lortnocoibon Asquith, andTrupanea dubautiae (Byran) (on the island of Kauai). The former two tephritids are deliberatelyintroduced biological control agents of weeds; and the latter two are endemic to the Hawaiian Islands. Ten species of hymenopterous parasitoids in six families were recovered from these four non-pest tephritids. Among these species of parasitoids, six were inadvertently introduced to Hawaii, includingEurytoma tephritidis Fullaway (Eurytomidae),Bracon terryi (Bridwell) (Braconidae),Habrocytus elevatus (Walker) (Pteromalidae),Euderus metallicus (Ashmead) (Eulophidae),Torymus advenus (Osten Sacken) (Torymidae), andEupelmus allynii (French) (Eupelmidae); and three were purposely introduced to Hawaii for classical biological control of economically important pests, includingEupelmus cushmani (Crawford),Diachasmimorpha tryoni (Cameron), (Braconidae), andDiachasmimorpha longicaudata (Ashmead). The relative abundance of parasitoid species and percent parasitism varied with the tephritid hosts. Our findings indicate that future development and implementation of biological control programs against frugivorous tephritids or other pests should consider the potential impact of some of the parasitoids on deliberately-introduced weed control agents. Before we can fully assess the potential impacts of some of the purposely-introduced parasitoids on non-target tephritids, however, effects of the many vagrant parasitoids associated with the non-targets should be thoroughly investigated     

The host- and microhabitat olfactory location by Fopius arisanus suggests a broad potential host range

Abstract.  The identification of infochemicals for parasitoid females is a critical issue in applied and fundamental parasitoid research. The olfactory location of host and its microhabitat by Fopius arisanus (Sonan, 1932) (Hymenoptera: Braconidae), an egg-pupal parasitoid of fruit flies (Diptera: Tephritidae), is investigated. Potential sources of volatiles are placed in opaque traps and tested in field cages, under seminatural conditions. Fopius arisanus females respond positively to synomones from mango leaves and from fruits belonging to many botanical families, including the nonhost plant strawberry. They prefer fruits infested by Tephritidae to uninfested ones but do not discriminate between fresh and old infestations. Fopius arisanus females are attracted by the odours of faeces of the tephritid fly Bactrocera zonata . They exhibit remote detection of a volatile kairomone coating the egg mass of all tested Tephritidae species but absent in the egg mass of the Muscidae Stomoxys calcitrans . All these infochemicals are volatile but only those emanating from fruit and from faeces are attractants perceived before landing. The relationships between this apparent generalist behaviour and the dietary specialization of F. arisanus are discussed, according to its ecology and behaviour in its natural environment.     

Impact of introduction of Bactrocera dorsalis (Diptera: Tephritidae) and classical biological control releases of Fopius arisanus (Hymenoptera: Braconidae) on economically important fruit flies in French Polynesia

Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), was discovered on Tahiti Island in July 1996. Eradication programs were conducted from 1997 to 2001, but failed. From 1998 to 2006, B. dorsalis was recovered from 29 different host fruit from the five Society Islands: Tahiti, Moorea, Raiatea, Tahaa, and Huahine. Analysis of coinfestation patterns by B. dorsalis, Bactrocera tryoni (Froggatt), and Bactrocera kirki (Froggatt) suggested B. dorsalis had displaced these two species and become the most abundant fruit fly in coastal areas. To suppress B. dorsalis populations, a classical biological control program was initiated to introduce the natural enemy Fopius arisanus (Sonan) (Hymenoptera: Braconidae) into French Polynesia from Hawaii. Wasps were released and established on Tahiti, Moorea, Raiatea, Tahaa, and Huahine Islands. In guava, Psidium guajava L., collections for Tahiti, F. arisanus parasitism of fruit flies was 2.1, 31.8, 37.5, and 51.9% for fruit collected for 2003, 2004, 2005 and 2006, respectively. Based on guava collections in 2002 (before releases) and 2006 (after releases), there was a subsequent decrease in numbers of B. dorsalis, B. tryoni, and B. kirki fruit flies emerging (per kilogram of fruit) by 75.6, 79.3, and 97.9%, respectively. These increases in F. arisanus parasitism and decreases in infestation were similar for other host fruit. Establishment of F. arisanus is the most successful example of classical biological control of fruit flies in the Pacific area outside of Hawaii and serves as a model for introduction into South America, Africa, and China where species of the B. dorsalis complex are established.     

Biology and host specificity of the chromolaena stem gall fly, Cecidochares connexa (Macquart) (Diptera: Tephritidae)

Abstract The stem-galling tephritid fly Cecidochares connexa (Macquart) was investigated as a potential biocontrol agent against the weed Chromolaena odorata (L) King & Robinson in Indonesia. Adults were tested in choice and no-choice tests, on 55 non-target plant species in 17 families. No oviposition was recorded on 53 of the species, while oviposition but no larval development was recorded on two. Field releases commenced in 1995 and establishment was immediate. The gall fly is now widely established in most Indonesian provinces, where gall parasitism and predation is generally low.     

Utilization of the Exotic Weed Pluchea odorata (Asteraceae) and Related Plants by the Introduced Biological Control Agent Acinia picturata (Diptera: Tephritidae) in Hawaii

The flower-head feeding fly Acinia picturata (Diptera: Tephritidae) was deliberately introduced from Mexico into Hawaii in 1959 for biological control of the exotic weed Pluchea odorata (Snow) (Asteraceae). Neither field efficacy nor possible non-target effects of the fly have been evaluated in the 40 years since the introduction. We assessed the impact of the fly on both its target host and on seven non-target plant species. The impact on the target weed was limited, with only 5-13% of the developing seeds in P. odorata flowerheads being destroyed by larval feeding. We did not detect any host range expansion of A. picturata onto flowerheads of two exotic or 5 endemic non-target plant species in the family Asteraceae.     

A compound produced by fruigivorous Tephritidae (Diptera) larvae promotes oviposition behavior by the biological control agent Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

Tephritid fruit fly parasitoids use fruit-derived chemical cues and the vibrations that result from larval movements to locate hosts sequestered inside fruit. However, compounds produced by the larvae themselves have not been previously described nor their significance to parasitoid foraging determined. We collected the volatiles from four species of tropical and subtropical Tephritidae: Anastrepha suspensa (Loew), Bactrocera dorsalis Hendel, Bactrocera cucurbitae Coquillett, and Ceratitis capitata (Wiedemann), representing two subfamilies (Dacinae and Trypetinae). Para-ethylacetophenone, an analog of a known tephritid parasitoid attractant, was a major constituent of all four, and was not associated with larvae of another acalypterate fly, Drosophila melanogaster Meigen, or with the calypterate Musca domestica L. It also was present in volatiles from whole, A. suspensa infested fruits of Eugenia uniflora (L.). Para-ethylacetophenone was not necessarily produced as a direct consequence of fruit consumption because it also was detected from larvae that developed in two artificial diets and in spent diets subsequent to larval development. Sensillae on both the antennae and ovipositor of the opiine braconid fruit fly parasitoid, Diachasmimorpha longicaudata (Ashmead) responded to the para-ethylacetophenone in larval volatiles and as a synthetic. Although a potential cue to foraging parasitoids, para-ethylacetophenone showed no long range (>1m) attractiveness to the adult female parasitoid, but did stimulate ovipositor-insertion and oviposition into both a natural (fruit) and an artificial (parafilm) substrate. Thus it may prove useful in colonizing and mass-rearing opine fruit fly parasitoids.     

Non-target host risk assessment of the idiobiont parasitoid Bracon celer (Hymenoptera: Braconidae) for biological control of olive fruit fly in California

Non-target risk posed by an African parasitoid, Bracon celer Szépligeti (Hymenoptera: Braconidae), was assessed for a classical biological control program against olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae: Dacinae), in California, USA. Behavioral and reproductive responses to non-target tephritid species were tested with beneficial (Chaetorellia succinea [Costa] and Parafreutreta regalis Munro) (Tephritidae: Tephritinae) and native (Rhagoletis fausta [Osten Sacken]) (Tephritidae: Trypetinae) fruit fly species in successive no-choice and choice experiments under close confinement in quarantine. Non-target host-plant substrates exposed to B. celer were yellow-starthistle flower heads containing C. succinea, Cape ivy stem galls containing P. regalis, and bitter-cherry fruit containing R. fausta. The parasitoid probed all three infested non-target plant substrates, but significantly less than olives infested with B. oleae. It produced offspring from P. regalis in Cape ivy stem galls, but appeared unable to penetrate yellow-starthistle flower heads with its ovipositor. Bracon celer killed some B. oleae and R. fausta larvae without parasitism. Reproduction on P. regalis indicates that B. celer has a broad physiological host range, which, combined with the parasitoid's acceptance of all three host-plant substrates, indicates a strong potential to negatively impact non-target species. Although physical and temporal barriers to host attack may reduce risk to most non-target tephritids by B. celer in California, the parasitoid should not be released due to its risk of harming the beneficial P. regalis. Release of P. regalis is still under consideration, however, and final risk assessment should depend on whether the fly proves useful for weed control.     

Taxonomy, attack strategies and host relations in flowerhead Tephritidae: a review

Abstract. 1. Host relations of Palaearctic flowerhead-inhabiting tephritids (Diptera: Tephritidae) are reviewed in terms of the interaction between type of attack strategy and flowerhead size.
2. Three types of attack strategy are recognized, each associated with a particular phase of Compositae flowerhead development. These strategies are characteristic of a tribe or subfamily, viz the Tephritini (Tephritinae), Terelliini (Tephritinae) and Myopitinae.
3. Tephritini species show the most generalized feeding method, and this enables them to attack both large and very small flowerheads. In contrast, the more specialized requirements of members of the Terelliini and Myopitinae apparently restricts these groups to plant taxa with large flowerheads. As a result, over the whole range of Compositae flowerhead sizes, there exists a positive relationship between flowerhead size and the diversity of associated tephritids.
4. The occurrence of stem-feeding tephritids, and differences between a coevolutionary and functional (colonization) explanation for tephritid host-relations are discussed.