Sunday Ekesi

Field infestation, life history and demographic parameters of the fruit fly Bactrocera invadens (Diptera: Tephritidae) in Africa

Field infestation rates of an invasive fruit fly species, Bactrocera invadens Drew Tsuruta & White on mango was determined at different localities in Kenya. At most of the locations and especially at low elevations, B. invadens frequently shared the same fruit with the indigenous fruit fly species Ceratitis cosyra (Walker) but often occurred at higher numbers than C. cosyra. The level of infestation varied with location ranging from 3.0 to 97.2 flies per kg of fruit. There was a significant inverse relationship between numbers of flies per kg of fruit and elevation at which fruit was collected, suggesting that B. invadens is a predominantly lowland pest. On an artificial diet, development of B. invadens immatures lasted 25 days; egg incubation required 1.2 days, larval development 11.1 days and puparia-adult development 12.4 days. About 55% of eggs developed to the adult stage. Life expectancy at pupal eclosion was 75.1 days in females and 86.4 days in males. Average net fecundity and net fertility were 794.6 and 608.1 eggs, respectively, while average daily oviposition was 18.2 eggs. Daily population increase was 11% and mean generation time was 31 days. Results are discussed in relation to the biology and ecology of the insect and in the development of mass rearing and control measures for B. invadens.

Host Plants and Host Plant Preference Studies for Bactrocera invadens (Diptera: Tephritidae) in Kenya, a New Invasive Fruit Fly Species in Africa

Abstract Bactrocera invadens Drew, Tsuruta & White (Diptera: Tephritidae), an invasive fruit fly species of Asian origin, was detected in Kenya in 2003, and is now well established in several parts of the country. We assessed the host range of this major quarantine pest in Kenya by collecting a wide range of cultivated and wild host plants from December 2004 to April 2006. Fruit were collected from 90 plant species representing 40 families from the Coast, Eastern, and Rift Valley provinces of the country where the fly population had been observed to occur in large numbers and where fruit and vegetable production is predominant. Fourteen plant species, among them cultivated and wild fruiting species, were found to be hosts of B. invadens. Fruit of mango, Mangifera indica L. (Anacardiaceae); banana Musa sp. AAA (Musaceae); and citrus [Citrus limon (L.) Burm.f. (lemon), Citrus reticulata Blanco (tangerine), and Citrus sinensis (sweet orange) (all Rutaceae)], were among the cultivated species heavily infested by B. invadens. Marula Sclerocarya birrea (A.Rich) Hochst. (Anacardiaceae) and Terminalia catappa L. (Combretaceae) were found to be the most infested noncultivated plants. These wild plants evidently ensure that sufficient reproductive bases exist for B. invadens during the off-season when the cultivated hosts are not in fruiting. In laboratory host preference studies, mango and banana were found to be the most preferred host plants among the nine cultivated plant species tested.

Evidence for Competitive Displacement of Ceratitis cosyra by the Invasive Fruit Fly Bactrocera invadens (Diptera: Tephritidae) on Mango and Mechanisms Contributing to the Displacement

ABSTRACT Bactrocera invadens Drew, Tsuruta & White (Diptera: Tephritidae) invaded Kenya in 2003. Before the arrival of B. invadens, the indigenous fruit fly species Ceratitis cosyra (Walker) was the predominant fruit fly pest of mango (Mangifera indica L.). Within 4 yr of invasion, B. invadens has displaced C. cosyra and has become the predominant fruit fly pest of mango, constituting 98 and 88% of the total population in traps and mango fruit at Nguruman, respectively. We tested two possible mechanisms responsible for the displacement namely; resource competition by larvae within mango fruit and aggression between adult flies. Under interspecific competition, larval duration in B. invadens was significantly shorter (6.2 ± 0.6-7.3 ± 0.3 d) compared with C. cosyra (8.0 ± 1.2-9.4 ± 0.4 d). Pupal mass in C. cosyra was affected by competition and was significantly reduced (7.4 ± 0.3-9.6 ± 0.6 mg) under competitive interaction compared with the controls (12.1 ± 1.5-12.8 ± 1.1 mg). Interspecific competition also had a significant adverse effect on C. cosyra eclosion, with fewer adults emerging under co-infestation compared with the controls. Interference competition through aggressive behavior showed that fewer C. cosyra (3.1 ± 0.8) landed on mango dome compared with the controls (14.2 ± 1.5) when adults were mixed with B. invadens adults in Plexiglas cages. Similarly the number of times C. cosyra was observed ovipositing was significantly lower (0.2 ± 0.2) under competitive interaction compared with the controls (6.1 ± 1.8). Aggressive encounters in the form of lunging/head-butting and chasing off other species from the mango dome was higher for B. invadens compared with C. cosyra. Our results suggest that exploitative competition through larval scrambling for resources and interference competition through aggressive behaviors of the invader are important mechanisms contributing to the displacement of C. cosyra by B. invadens in mango agroecosystems.

Mortality in Three African Tephritid Fruit Fly Puparia and Adults Caused by the Entomopathogenic Fungi, Metarhizium anisopliae and Beauveria bassiana

The pathogenicity of 13 isolates of Metarhizium anisopliae and two isolates of Beauveria bassiana to Ceratitis capitata and Ceratitis var. rosa fasciventris exposed as late third instar larvae in sand was evaluated in the laboratory. All isolates caused a significant reduction in adult emergence and a corresponding large mortality on puparia of both species. All isolates also induced large deferred mortality in emerging adults following treatment as late third instar larvae. On C. capitata , seven isolates ( M. anisopliae ICIPE 18, 20, 32, 60 and 69 and B. bassiana ICIPE 44 and 82) caused significantly higher mortality on puparia than other isolates. With the exception of ICIPE 32, the other four isolates of M. anisopliae above were the most pathogenic against C. r. fasciventris . Dose-response study carried out with these isolates of M. anisopliae on the two species of flies above plus another species, Ceratitis cosyra showed that the dose-mortality regression lines of ICIPE 18 and 20 were steeper with lower LC 50 values when compared with ICIPE 60 and 69 on the three species. When these two isolates were evaluated with regard to their pathogenicity to different pupal age, adult emergence was found to increase with increasing pupal age with a corresponding decrease in mortality in puparia and emerging adults in the three species of fruit flies. M. anisopliae ICIPE 18 and 20 were equally pathogenic to all pupal ages tested in C. capitata and C. cosyra but ICIPE 18 was more pathogenic to older puparia of C. r. fasciventris than ICIPE 20. Our results suggest that soil inoculation with M. anisopliae under mango trees might form an important component of integrated pest management strategies in areas where these three species of fruit fly coexist.

Ecological niche and potential geographic distribution of the invasive fruit fly Bactrocera invadens (Diptera, Tephritidae)

Two correlative approaches to the challenge of ecological niche modeling (genetic algorithm, maximum entropy) were used to estimate the potential global distribution of the invasive fruit fly, Bactrocera invadens, based on associations between known occurrence records and a set of environmental predictor variables. The two models yielded similar estimates, largely corresponding to Equatorial climate classes with high levels of precipitation. The maximum entropy approach was somewhat more conservative in its evaluation of suitability, depending on thresholds for presence/absence that are selected, largely excluding areas with distinct dry seasons; the genetic algorithm models, in contrast, indicate that climate class as partly suitable. Predictive tests based on independent distributional data indicate that model predictions are quite robust. Field observations in Benin and Tanzania confirm relationships between seasonal occurrences of this species and humidity and temperature.

Pathogenicity of Metarhizium anisopliae (Metsch.) Sorokin and Beauveria bassiana (Balsamo) Vuillemin, to three adult fruit fly species: Ceratitis capitata (Weidemann), C. rosa var. fasciventris Karsch and C. cosyra (Walker) (Diptera :Tephritidae)

The pathogenicity of two isolates of Beauveria bassiana and 12 of Metarhizium anisopliae towards adult fruit flies, Ceratitis capitata and Ceratitis rosa var. fasciventris was tested in the laboratory. Fruit flies were exposed to dry conidia evenly spread on velvet material covering the inner side of a cylindrical plastic tube. All isolates tested were pathogenic to both species of fruit flies. Mortality ranged from 7 to 100% in C. capitata and from 11.4 to 100% in C. rosa var. fasciventris at 4 days post-inoculation. Six isolates, M. anisopliae ICIPE 18, 20, 32, 40, 41 and 62, were highly pathogenic to both C. capitata and C. rosa var. fasciventris. The LT90 values of the most pathogenic isolates ranged between 3–4 days in both insects. Because of the difficulties in rearing C. cosyra, only the isolates that were highly pathogenic to both C. rosa var. fasciventris and C. capitata were tested against adult C. cosyra. They caused mortality of between 72–78% at 4 days post-inoculation. The LT90 values in all the isolates did not exceed 4 days. One of the most pathogenic isolates, M. anisopliae ICIPE 20, was evaluated against C. capitata and C. rosa var. fasciventris in cage experiments using threeautoinoculators (maize cob, cheesecloth and Petri dish) in an autoinoculative deviceconsisting of plastic mineral bottle. Mortality of between 70–93% was observed in flies of both species that were captured from the cages and held under laboratoryconditions. These results indicate the possibility of fruit fly suppression withentomopathogenic fungi using an autoinoculative device.

Effect of Temperature on Germination, Radial Growth and Virulence of Metarhizium anisopliae and Beauveria bassiana on Megalurothrips sjostedti

The effect of temperature on germination, radial growth and pathogenic activity of two strains of Beauveria bassiana and four strains of Metarhizium anisopliae on the legume flower thrips, Megalurothrips sjostedti , was studied in the laboratory. Germination, radial growth and pathogenic activity was low for all strains at 15 C. Optimum temperature for germination, radial growth and pathogenic activity ranged between 25-30 C. The fastest growing strain at 25-30 C was M. anisopliae strain ICIPE 69. Compared to other strains, ICIPE 69 also appeared to have a broad temperature range of pathogenic activity against the pest. The significance of these findings in relation to control of M. sjostedti is discussed.

Effect of soil temperature and moisture on survival and infectivity of Metarhizium anisopliae to four tephritid fruit fly puparia.

The infectivity of 4 isolates of Metarhizium anisopliae to puparia of Ceratitis capitata treated as late third-instar larvae in unsterilized soil was investigated in the laboratory under controlled temperature and moisture. At 20-30 degrees C, mortality in puparia was highest at water potential of -0.1 and -0.01 mega Pascal (MPa) and lowest at water potential of -0.0055 and -0.0035 MPa in all the isolates. In wetter soil however, isolates ICIPE 20 and 60 caused significantly higher mortality than ICIPE 18 and 69. The survival of conidia in drier soil (-0.1 MPa) was not adversely affected at all temperatures. However, in wet soil (-0.0035 MPa) there was drastic reduction in colony counts in ICIPE 18 and 69 at 25 and 30 degrees C but conidial density in ICIPE 20 and 60 remained at the initial level at 14 days after inoculation at all temperatures. When ICIPE 20 was evaluated against three other fruit fly species (Ceratitis cosyra, Ceratitis rosa, and Ceratitis fasciventris), significant reduction in adult emergence and higher pupal mortality occurred in C. cosyra and C. fasciventris than in C. rosa at a combination of 15 and 20 degrees C and -0.1 and -0.0035 MPa. However, at higher temperature and the same moisture level, the isolates were equally pathogenic across the 3 species. It is probable that in addition to pathogen cycling and multiplication from dead infected insects in the soil, a balance between microbial degradation and replenishment of inoculum of virulent isolates occur through fluctuations in, and intricate interactions between temperature and moisture levels. This study is indicative of the potential of using isolate ICIPE 20 for soil inoculation against pupariating third-instar larva of fruit flies, thus providing a novel alternative to chemical soil application.

Susceptibility of Megalurothrips sjostedti developmental stages to Metarhizium anisopliae and the effects of infection on feeding, adult fecundity, egg fertility and longevity

The susceptibility of immature stages of the legume flower thrips, Megalurothrips sjostedti, to the entomopathogenic fungus Metarhizium anisopliae, was investigated under laboratory conditions. The adult stage was found to be more susceptible to infection than the larval and pupal stages. Mortality at all stages was dose‐dependent, with the highest concentration of 1×108 conidia ml−1 producing the highest mortality (26, 46 and 100% for larvae, pupae and adults, respectively) at 8 days post‐inoculation. At the same concentration, daily pollen consumption was significantly reduced at 2 days after treatment in infected adults but more slowly in infected larvae. Fecundity, egg fertility and longevity in adults surviving infection as larvae were significantly reduced compared to the control.

Adaptation to and Small-Scale Rearing of Invasive Fruit Fly Bactrocera invadens (Diptera: Tephritidae) on Artificial Diet

Abstract Larval rearing of Bactrocera invadens Drew, Tsuruta & White (Diptera: Tephritidae) on artificial diet is described. The adaptation process for this insect, when moved from whole mango, Mangifera indica L., fruit rearing to artificial diet based on wheat bran, took between three and five generations to reach the plateau of quality control parameters observed for rearing the insect on whole mango fruit. Small-scale rearing on wheat, Triticum aestivum L.-, or carrot, Daucus corata-based diet revealed significantly higher pupal recovery for flies reared on the wheat-based artificial diet (68.8%) compared with the carrot-based diet (58.2%). Weekly production of puparia was 3,966.8 on wheat- and 3,012.1 on carrot-based diet. Other quality control parameters, including pupal weight, adult emergence, flight ability, fecundity, and fertility did not differ significantly between the two artificial rearing media tested.