Foraging Behavior and Patch Time Allocation by Fopius arisanus (Hymenoptera: Braconidae), an Egg-Larval Parasitoid of Tephritid Fruit Flies

This study quantitatively describes the host-searching behavior of Fopius arisanus (Sonan) (Hymenoptera: Braconidae), an important egg-larval parasitoid of tephritid fruit fly pests, on coffee berries infested with host eggs of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). We also investigate the parasitoid's response to local variation in host patch quality. The temporal pattern of behavioral organisation was examined by constructing an ethogram. The parasitoid spent over 90% of its foraging time in detecting and locating hosts after arriving on a host-infested fruit, and displayed a relatively fixed behavioral pattern leading to oviposition. Patch residence time increased in the presence of host-associated cues, following successful ovipositions, and with increasing size of host clutches per fruit, but decreased with each successive visit to the same host patch and with increasing availability of alternative host patches. The parasitoid females discriminated against previously parasitized hosts and spent significantly less time and searching effort on patches previously exploited by herself or by conspecific females. The effective host-searching behavior, perfect host discrimination ability, and success-motivated searching strategy shown by F. arisanus ensured a thorough exploitation of host resources by this parasitoid.     

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.     

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.     

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.     

Evaluation of Beauveria bassiana and B. brongniartii strains and four wild-type fungal species against adults of Bactrocera oleae and Ceratitis capitata

The virulence of two isolates of the hyphomycete fungi, Beauveria bassianaand B. brongniartii, and additional fungal species isolated from diseased Bactrocera oleae pupae and Sesamia nonagrioideslarvae were assessed against adults of the olive fruit fly B. oleae and the Mediterranean fruit fly Ceratitis capitata (Diptera: Tephritidae). Contact and oral bioassays revealed that moderate to high mortality rates for the olive fruit fly occurred when the adults were exposed to conidia of Mucor hiemalis, Penicillium aurantiogriseum, P. chrysogenum and B. bassianaisolates. A strain of M. hiemalis isolated from S. nonagrioides larvae was the most toxic resulting in 85.2% mortality to the olive fruit fly adults. B. brongniartiiand B. bassiana were the most pathogenic to the C. capitataadults causing 97.4 and 85.6% mortality. Metabolites collected from the M. hiemalis and P. chrysogenum isolates were toxic to adults of both 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.     

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.     

Mass‐rearing optimization of the parasitoid Psyttalia lounsburyi for biological control of the olive fruit fly

The olive fruit fly, Bactrocera oleae (Tephritidae), is a direct pest of olives that has invaded the Mediterranean Region and California. Psyttalia lounsburyi (Braconidae), a larval parasitoid from Africa, has been approved for release in the USA as a classical biological agent. However, it has been difficult to rear the parasitoid in the laboratory because it is multivoltine, and the host develops only in fresh olives, which are not available for most of the year. A method to rear the parasitoid on the factitious host, Mediterranean fruit fly (Ceratitis capitata) was developed, but it was not very efficient for producing large numbers of parasitoids needed for release. We developed a number of ways to improve the efficiency of rearing, including the frequency and duration of exposure for oviposition, optimizing the density of adult parasitoids, host age, as well as methods to quickly standardize the number of larvae exposed and to count emerging adult parasitoids. We significantly improved the number of progeny produced per female and the sex ratio of progeny. Thanks to these improvements, we produced in 2017 over 119,000 adults and shipped over 53,900 for release in California.     

Cross mating studies among five fruit fly parasitoid populations: potential biological control implications for tephritid pests

The reproductive compatibility between four different species/populations of the tephritid parasitoid Psyttalia (Walker) species from Kenya and individuals of the morphologically identical Psyttalia concolor (Szépligeti) (Hymenoptera: Braconidae) from a laboratory culture in Italy used in augmentative biological control of olive fly, Bactrocera oleae (Gmelin) (Diptera: Tephritidae) was assessed through cross mating tests using single-pair and group mating methods. Reciprocal crosses among the species resulted in the production of viable offsprings up to the second generation. In spite of the successful production of viable offspring in the laboratory, Psyttalia species are known to have specific host fruit and/or host fly preferences and populations/species may be isolated in one way or the other. However, it is not known whether these populations/species interbreed in the field. We discuss the ability of these parasitoids to interbreed and the potential effects of that on their use as biological control agents, especially in environments where other closely related species are present or in situations where multiple parasitoid introductions are intended.     

<Emphasis Type="Italic">Fopius arisanus</Emphasis> oviposition in four <Emphasis Type="Italic">Anastrepha</Emphasis> fruit fly species of economic importance in Mexico

We studied the oviposition performance of Fopius arisanus (Sonan) (Hymenoptera: Braconidae) attacking eggs of four fruit flies of the genus Anastrepha Schiner (Diptera: Tephritidae) under laboratory conditions. The complete process of oviposition on an individual egg of Anastrepha ludens (Loew) lasts in average 85.4 ± 2.9 s, including a tremor (25.8 ± 1.03 s) observed in the middle of this process related to the egg’s descent. The average parasitism of A. ludens egg was 60.9 ± 7.5%, with only 1.2% of superparasitized eggs. During individual acts of oviposition, we noted that F. arisanus possesses a highly flexible ovipositor that curves easily as it searches for additional suitable eggs, which may be of particular benefit when a female finds large clutches of eggs. The individual oviposition of F. arisanus in host fruits attacked by Anastrepha spp. varies with the egg clutch size of each fruit fly species: A. serpentina laid the biggest egg clutches (21.3 ± 1.4), followed by A. ludens (14.2 ± 0.9), and A. striata (1.0 ± 0.0) (=A. obliqua). The time spent by F. arisanus in individual ovipositions was parallel to these findings, reinforcing the idea that F. arisanus attacks several eggs in each individual insertion of its ovipositor. Neither formal oviposition acts, nor adult emergences of F. arisanus were registered in A. obliqua. We discuss the potential of F. arisanus as natural enemy of fruit flies of the genus Anastrepha, and explore the eventual developing of its mass rearing. Handling Editor: Torsten Meiners.     

The biology of Bracon celer as a parasitoid of the olive fruit fly

A series of laboratory experiments was conducted on a colony of Bracon celer Szépligeti (Hymenoptera: Braconidae) reared on the olive fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae). Female B. celer preferentially probe and oviposit into olives containing late third-instar fly larvae. The parasitoid develops as a solitary, ectoparasitic idiobiont. Mean development time (oviposition to adult eclosion) at 22 °C was, for females, 36±1 (SE) days, and for males, 34±1 days. The mean longevity of adult female wasps when provided honey and water was significantly greater than when they were provided water alone, or nothing. The females produced an average of 9.7±7.2 progeny during their lifetimes, but production levels in the insectary colony suggested that this level of fecundity was artificially low and could be improved. The discrepancy may be a consequence of constraints on oviposition behavior imposed by the experimental design. The results are discussed with respect to insectary production methods and the potential use of B. celer as a biological control agent for olive fly in California.     

Evaluation of Fopius arisanus as a biological control agent for the olive fruit fly in California

1 The egg‐prepupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) was evaluated in quarantine facilities as a potential biological control agent for the olive fruit fly Bactrocera oleae (Diptera: Tephritidae) in California, U.S.A. 2 Nontarget testing of two weed biological control agents confirmed that F. arisanus will not attack Tephritidae that feed in inflorescences or galls. It may, however, pose risks to native Tephritidae that feed in fruit. 3 Females preferentially oviposited in eggs, although first‐instar B. oleae were also attacked. Low lifetime reproductive potential and high rates of direct mortality inflicted on host eggs indicate that rearing on B. oleae may prove difficult. 4 In multiparasitized B. oleae, F. arisanus prevailed in competition against two species of larval–pupal parasitoids, Diachasmimorpha kraussii and Psyttalia concolor (both Hymenoptera: Braconidae). 5 The broad host‐range of F. arisanus with respect to fruit‐feeding Tephritidae may preclude its introduction to California, as may its low fecundity and its intrinsic competitive superiority over larva l–pupal parasitoids, which include specialists on B. oleae that are currently being introduced to California. High rates of direct mortality, however, point to potential uses in augmentative biological control. Whether or not F. arisanus is released in California, its biology as a parasitoid of B. oleae has been little studied to date and the results herein may be applied in other regions worldwide where B. oleae is a problem.     

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.     

Old and new host-parasitoid associations: parasitism of the invasive fruit fly Bactrocera invadens (Diptera: Tephritidae) and five African fruit fly species by Fopius arisanus,an Asian opiine parasitoid

Fopius arisanus (Sonan), a solitary koinobiont endoparasitoid of fruit flies, was introduced for testing and final release against the recently discovered species Bactrocera invadens Drew, Tsuruta and White in Africa. Laboratory experiments were conducted to assess host preference, host acceptability for oviposition, and physiological suitability of B. invadens and five other indigenous tephritid fruit fly species, namely, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), C. cosyra (Walker), C. rosa Karsch, C. fasciventris (Bezzi), and C. anonae Graham. Females of F. arisanus visited all host egg patches, but showed a stronger preference to eggs of B. invadens, which was also most accepted for oviposition. Successful development of parasitoid progenies varied greatly across hosts, with B. invadens yielding the highest parasitoid progeny and C. fasciventris yielding no F. arisanus progeny. Most of the parasitoid eggs laid in C. rosa and C. fasciventris were encapsulated. Sex ratio was not influenced by host species and it was female biased in all hosts that produced parasitoid progeny. Fopius arisanus was able to establish a new association with C. capitata, C. cosyra and to a lesser extent C. anonae. The results are discussed in the light of the potential use of F. arisanus as a biological control agent of B. invadens.     

Host specificity of the egg pupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) in La Reunion

Fopius arisanus is a polyphagous parasitoid of Tephritidae, which has been recently introduced to La Réunion Island as part of a classical biological control programme. We carried out laboratory experiments to assess the host specificity of this parasitoid, initially reared on Bactrocera zonata, and then offered for parasitization the eight local tephritid pest species. Naive or experienced parasitoid females were given tephritid eggs in no choice tests. Fopius arisanus females parasitize all fly species but parasitism varies with host species. No adult wasps emerge from Bactrocera cucurbitae and the survival of this species is only slightly affected by parasitism. Dissections show that the late instars of this fly may eliminate the parasitoid by encapsulation. When developing on Ceratitis capitata, Ceratitis rosa, Dacus ciliatus, Dacus demmerezi, and Neoceratitis cyanescens, parasitoid survival rate ranges from 10 to 25%. Bactrocera zonata and Ceratitis catoirii are the best hosts, yielding parasitoid survival rates of more than 70% with no premature mortality. The egg-larval mortality of C. capitata, C. rosa, D. ciliatus, and N. cyanescens, and the pupal mortality of D. demmerezi, are significantly increased by parasitism. The size of emerging adults is affected by host species and is correlated to pupal weight. Bactrocera zonata would be a favorable host to support routine colonization of F. arisanus for mass production of this parasitoid.     

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.     

Comparative evaluation of two olive fruit fly parasitoids under varying abiotic conditions

Psyttalia lounsburyi (Silvestri) and P. humilis (Silvestri) (Hymenoptera: Braconidae) were evaluated in California for their potential to control the invasive olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae). Psyttalia lounsburyi is a specialist on B. oleae while P. humilis also attacks other tephritid species. Field cage trials, conducted from 2006 to 2009, were used to compare P. lounsburyi and two populations of P. humilis (Kenya and Namibia) in California’s interior valley and coastal regions. Both parasitoid species reproduced on B. oleae in all trials. Under similar abiotic conditions, offspring production per female was higher in P. humilis than in P. lounsburyi, suggesting that host specificity by P. lounsburyi does not confer a higher efficiency on B. oleae in cultivated olives. Two abiotic factors were shown to impact parasitoid efficiency. First, adult parasitoid survival was poor during periods of high summer temperatures, common to the olive production areas in California’s interior valleys. Second, parasitism levels were lower on B. oleae larvae feeding in larger Ascolano cv. fruit than in smaller Manzanillo cv. fruit. Results are discussed relative to biological control of B. oleae in commercial olives and the usefulness of natural enemies specialized to attack fruit flies in wild olives compared with the larger cultivated olive fruit.     

Crop domestication relaxes both top-down and bottom-up effects on a specialist herbivore

Domestication of crop plants selects for numerous traits that often distinguish them dramatically from their wild progenitors. In some cases, these modifications lead to increased herbivory, by enhancing their attractiveness to herbivorous insects or reducing the efficiency of natural enemies, or both. This study investigated the effects of fruit enlargement on the olive (Olea europaea L.), the specialist olive fruit fly, Bactrocera oleae (Rossi), and its specialized larval parasitoids. Wild olive fruit are small (<2 mm pulp thickness) and the larval parasitoids associated with B. oleae have short ovipositors (<3 mm), while cultivated fruit are larger (4–8 mm pulp thickness). Female flies allocate more offspring to large than to small fruit within or across different-sized commercial cultivars, without reducing the fitness of their offspring. Fly larvae move deeper into the olive pulp with their increasing age and fruit size. In contrast, the specialist larval parasitoid, Psyttalia lounsburyi (Silvestri), more effectively parasitizes hosts in smaller than larger fruit. The inverse relationship between the performance of the fly and its co-evolved parasitoids on fruit of increasing sizes indicates that olive cultivation favors the success of the fly by providing a better food resource and more enemy-free space. These findings offer some explanation for the failure of the decades-old classical biological efforts to manage B. oleae using specialized larval parasitoids in the Mediterranean Basin and provide further evidence that crop domestication can alter host–parasitoid interactions.     

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.     

Flower nitrogen status and populations of Frankliniella occidentalis feeding on Lycopersicon esculentum

Alighting and capture of wild olive fruit flies, Bactrocera oleae (Rossi) (Diptera, Tephritidae), on spheres of seven different colours was studied on Chios island, Greece. The 70-mm-diam plastic spheres, coated with adhesive, were suspended on olive trees. Yellow and orange spheres trapped the greatest number of males while red and black spheres trapped the greatest number of females. White and blue spheres were the least effective for both sexes. Peak captures occurred in the late afternoon and especially around sunset. Since mating takes place in the last hours of the photophase, the increased captures during that period may be related to the sexual behaviour of the fly. When red spheres were assessed against glass McPhail traps baited with 2% ammonium sulphate, which consist a standard tool for monitoring the olive fruit fly in Greece, there were no significant differences in male captures. However, spheres trapped almost three times as many females as McPhail traps. The possible mechanisms underlying colour discrimination, the motivation of alighting flies and the possible use of red spheres for monitoring and controlling B. oleae are discussed.