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.     

Acceptability and suitability of six fruit fly species (Diptera: Tephritidae) for Kenyan strains of Psyttalia concolor (Hymenoptera: Braconidae)

Host acceptability and suitability Psyttalia concolor (Szépligeti) is a koinobiont, larval parasitoid of tephritid fruit flies. Individuals of P. concolor were field-collected from coffee in the central highlands of Kenya, and cultured initially on Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann). They were then examined for their ability to oviposit in and develop on five other tephritid species that are pests in Kenya. In addition to the medfly, acceptability for oviposition and suitability for development were tested against the mango fruit fly, Ceratitis cosyra (Walker), the Natal fruit fly, Ceratitis rosa Karsch, Ceratitis fasciventris (Bezzi), Ceratitis anonae Graham and the melon fruit fly, Bactrocera cucurbitae (Coquillett). Ceratitis capitata and C. cosyra were accepted as hosts significantly more often than the other species. Superparasitism was recorded only from C. capitata and C. cosyra. Two days after oviposition, parasitoid eggs in C. fasciventris and B. cucurbitae were encapsulated, whereas those in C. rosa and C. anonae were encapsulated, and often melanized. Ceratitis capitata was the most suitable host for Kenyan populations of Psyttalia concolor in terms of progeny production, and proportion of female progeny.     

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

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

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.     

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.     

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

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

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.     

Effect of constant temperatures on germination, radial growth and virulence of Metarhizium anisopliae to three species of African tephritid fruit flies

The effect of temperatureon conidial germination, mycelial growth, andsusceptibility of adults of three tephritidfruit flies, Ceratitis capitata(Wiedemann), C. fasciventris (Bezzi) andC. cosyra (Walker) to six isolatesof Metarhizium anisopliae were studied inthe laboratory. There were significantdifferences among the isolates in the effect oftemperature on both germination and growth.Over 80% of conidia germinated at 20, 25 and30°C, while between 26 and 67% conidiagerminated at 35°C and less than 10% at15°C within 24 hours. Radial growth was slowat 15°C and 35°C with all of theisolates. The optimum temperature forgermination and mycelial growth was 25°C. Mortality caused by the six fungal isolatesagainst the three fruit fly species varied withtemperature, isolate, and fruit fly species.Fungal isolates were more effective at 25, 30and 35°C than at 20°C. The LT₉₀values decreased with increasing temperature upto the optimum temperature of 30°C. Therewere significant differences in susceptibilitybetween fly species to fungal infection at allthe temperatures tested.     

Fruit fly liquid larval diet technology transfer and update

Since October 2006, the US Department of Agriculture–Agricultural Research Service (USDA–ARS) has been implementing a fruit fly liquid larval diet technology transfer, which has proceeded according to the following steps: (1) recruitment of interested groups through request; (2) establishment of the Material Transfer Agreement with agricultural research service; (3) fruit fly liquid larval diet starter kit sent to the requestor for preliminary evaluation; (4) problem‐solving through email or onsite demonstration; (5) assessment on feedback from the participants to decide whether to continue the project. Up to date, the project has involved 35 participants from 29 countries and 26 species of fruit flies. Fourteen participants have concluded their evaluation of the process, and 11 of these 14, have deemed it to be successful. One participant has decided to implement the project on a larger scale. The 14 participants were, Argentina (Ceratitis capitata and Anastrepha fraterculus), Bangladesh (Bactrocera cucurbitae, C. capitata, and Bactrocera dorsalis), China (Fujia province) (B. dorsalis), Italy (C. capitata), Fiji (Bactrocera passiflorae), Kenya (Bactrocera invadens, Ceratitis cosyra), Mauritius (Bactrocera zonata and B. cucurbitae), Mexico (Anastrepha species), Philippines (Bactrocera philippinese), Thailand (Bactrocera correcta), Austria (C. capitata, Vienna 8 and A. fraterculus), Israel (Dacus ciliatus and C. capitata), South Africa (C. capitata, Vienna 8) and Australia (C. capitata). The Stellenbosch medfly mass‐rearing facility in South Africa and the CDFA in Hawaii were two mass‐scale rearing facilities that allowed us to demonstrate onsite rearing in a larger scale. Demonstrations were performed in CDFA in 2007, and in Stellenbosch, South Africa in 2008; both were found to be successful. The Stellenbosch medfly mass‐rearing facility in South Africa decided to adopt the technology and is currently evaluating the quality control of the flies that were reared as larvae on a liquid diet.     

Response of two parasitoid species (Hymenoptera: Braconidae,Figitidae) to tephritid host and host food substrate cues

The Neotropical‐native figitid Aganaspis pelleranoi (Brèthes) and the Asian braconid Diachasmimorpha longicaudata (Ashmead) are two parasitoids of Tephritidae fruit flies with long and recent, respectively, evolutionary histories in the Neotropics. Both species experienced a recent range of overlap. In Argentina, A. pelleranoi is a potential species in biological control programs against the pestiferous tephritid species, Anastrepha fraterculus (Wiedemann) and Ceratitis capitata (Wiedemann), whereas D. longicaudata is already used in open‐field releases against Medfly in central‐western Argentina. To characterize the host‐foraging strategies of A. pelleranoi and D. longicaudata, olfactometer experiments were conducted comparing responses to C. capitata and A. fraterculus larvae, in two kinds of food substrate: fruit and artificial larval medium. To control the possible influence of host larvae used for parasitoid rearing on olfactory response, two strains of both parasitoid species, reared on both tephrtid species, were studied. Volatiles directly emanating either from A. fraterculus or C. capitata larvae may be detected by both A. pelleranoi and D. longicaudata, although chemical stimuli originating from the combination of host larvae and the habitat of the host were preferred. However, olfactory cues associated with host larvae probably play a relevant role in host searching behaviour of A. pelleranoi, whereas for D. longicaudata, the host‐habitat olfactory stimuli would be highly essential in short‐range host location. The strain of the parasitoids did not affect host search ability on the two tephritid species evaluated. These evidences are relevant for mass production of both parasitoids and their impact following open‐field augmentative releases.     

Odour-mediated foraging by yellowjacket wasps (Hymenoptera: Vespidae): predation on leks of pheromone-calling Mediterranean fruit fly males (Diptera: Tephritidae)

Predation is probably the most important male mortality factor in insect species with courtship displays that render males performing them conspicuous targets of predators. Sexually active Mediterranean fruit fly males, Ceratitis capitata (Wied.), aggregate in leks, where they participate in agonistic encounters and engage in visual, acoustic and pheromone-calling displays to attract receptive females. The objective of this study was to assess: a) whether sexually displaying C. capitata males in leks inside host and non-host foliage are subject to predation by the most prominent predators yellow-jacket wasps, Vespula germanica (F.), and if so, b) whether olfactory, visual or auditive stimuli are used by foraging wasps in locating male C. capitata prey. Studies were carried out in a citrus orchard and surroundings on the island of Chios, Greece. Observations were conducted using perforated containers hung within mulberry, fig or citrus foliage. Living C. capitata flies of different sex and either mature or immature were placed inside. Our results show that the yellowjacket wasps have learned to associate the presence of sexually active medfly males aggregated in leks with their prey's pheromone (kairomone). Foraging wasps, flying through the crowns of host trees, responded to the odour source of C. capitata male pheromone by approaching from downwind. Even inside dense citrus tree foliage, wasps keyed in on aggregations of pheromone-calling males using olfactory stimuli. Stimuli of visual and acoustic male signalling were only used at close range, after having followed the pheromone plume close to its source. Visual cues played a greater role in directing wasp foraging under more open and exposed host foliage conditions. Odour-based foraging of wasps inside host foliage in the mid-morning hours, when medfly male lekking activities peak, shifted gradually to a more visual-based host fruit patrolling in the afternoons to capture ovipositing and feeding medfly females. On ripe fruit, particularly fig, V. germanica visual prey hunting also included the capture of feeding medfly males, other feeding Diptera, as well as medfly larvae extracted from wasp-made perforations in the fruit.     

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.     

Conditional embryonic lethality to improve the sterile insect technique in Ceratitis capitata (Diptera: Tephritidae)

Background  

The sterile insect technique (SIT) is an environment-friendly method used in area-wide pest management of the Mediterranean fruit fly Ceratitis capitata (Wiedemann; Diptera: Tephritidae). Ionizing radiation used to generate reproductive sterility in the mass-reared populations before release leads to reduction of competitiveness.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

Ecological niches and potential geographical distributions of Mediterranean fruit fly (Ceratitis capitata) and Natal fruit fly (Ceratitis rosa)

Aim To predict and compare potential geographical distributions of the Mediterranean fruit fly (Ceratitis capitata) and Natal fruit fly (Ceratitis rosa). Location Africa, southern Europe, and worldwide. Methods Two correlative ecological niche modelling techniques, genetic algorithm for rule‐set prediction (GARP) and a technique based on principal components analysis (PCA), were used to predict distributions of the two fly species using distribution records and a set of environmental predictor variables. Results The two species appear to have broadly similar potential ranges in Africa and southern Europe, with much of sub‐Saharan Africa and Madagascar predicted as highly suitable. The drier regions of Africa (central and western regions of southern Africa and Sahelian zone) were identified as being less suitable for C. rosa than for C. capitata. Overall, the proportion of the region predicted to be highly suitable is larger for C. capitata than for C. rosa under both techniques, suggesting that C. capitata may be tolerant of a wider range of climatic conditions than C. rosa. Worldwide, tropical and subtropical regions are highlighted as highly suitable for both species. Differences in overlap of predictions from the two models for these species were observed. An evaluation using independent records from the adventive range for C. capitata and comparison with other predictions suggest that GARP models offer more accurate predictions than PCA models. Main conclusions This study suggests that these species have broadly similar potential distributions worldwide (based on climate), although the potential distribution appears to be broader for C. capitata than for C. rosa. Ceratitis capitata has become invasive throughout the world, whereas C. rosa has not, despite both species having broadly similar potential distributions. Further research into the biology of these species and their ability to overcome barriers is necessary to explain this difference, and to better understand invasion risk.     

Patterns of resource distribution among conspecific larvae in two fruit fly species: Anastrepha fraterculus and Ceratitis capitata (Diptera: Tephritidae)

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Attraction of female Chiracanthium mildei(Araneae: Clubionidae) to olfactory cues from maleMediterranean fruit flies Ceratitis capitata(Diptera: Tephritidae)

Chiracanthium mildei, a common sac spider (Clubionidae) in Israel, is an important predator of pest insects in citrus and apple orchards. Field observations indicate that the females of this hunting spider are attracted to traps containing the parapheromone trimedlure and males of the Mediterranean fruit fly (Ceratitis capitata),which were caught in these traps. Laboratory testsshow that C. mildei females are attractedto olfactory cues from male C. capitata, but notto trimedlure. This suggests that C. mildeimay be an important nocturnal predator of C.capitata.     

Effects of recent experience on foraging in tephritid fruit flies

In field-cage studies, we investigated how the foraging behavior of tephritid fruit flies is modified by experience immediately prior to release on host plants. We observed females of a relatively monophagous species,Rhagoletis mendax (blueberry maggot fly), an oligophagous species,Rhagoletis pomomella (apple maggot fly), and a polyphagous species,Ceratitis capitata (Mediterranean fruit fly). Just prior to release on a host plant, the following kinds of stimuli were supplied: (1) single oviposition in a host fruit, (2) contact with 20% sucrose, (3) contact with a mixture of protein food (bird feces and sucrose), (4) contact with water, and (5) a walk over a host-plant leaf. When flies foraged on host plants without resources, search was most intensive (as measured by number of leaves visited) following a single oviposition in fruit, but residence time generally was the same following exposure to sugar, protein, and fruit stimuli.Rhagoletis mendax andC. capitata females visited the fewest leaves following exposure to water or host leaves, whereasR. pomonella foraged equally intensively following exposure to food stimuli, water, or leaves. On host plants containing resources (fruit and protein food), a single oviposition dramatically increased the number of females of all three species that found fruit compared to females that received experience with food, water or foliar stimuli. We found no significant effect of recent brief experience with any of the stimuli on subsequent attraction to protein food. Overall,C. capitata exhibited a higher propensity to abandon host plants than eitherR. mendax orR. pomonella. We suggest that this may reflect adaptations to differences in distribution of host plants in nature, strategies of dispersal, and host range.