Slawomir A. Lux

A New Invasive Fruit Fly Species from the Bactrocera dorsalis (Hendel) Group Detected in East Africa

Abstract—A new fruit fly species suspected to be from the Bactrocera dorsalis (Hendel) group (originating from Asia), was detected during routine field surveys in the Coast Province of Kenya. Since most species in this group are of tremendous quarantine concern when introduced, and considering the fact that it has never before been detected or reported in continental Africa, surveys were immediately initiated covering a distance of over 3000 km across major fruit-growing and trading localities within Kenya, to determine the extent of spread of the new invasive species. We report on the detection of the flies, preliminary results of the survey, and discuss the potential effects of these flies on the horticulture industry in East Africa.Résumé—Une nouvelle mouche des fruits appartenant vraisemblablement au groupe Bactrocera dorsalis (Hendel) (originaire dTnde), a été trouvée lors d’enquêtes de terrain de routine dans la province côtière du Kenya. Depuis que la plupart des espèces de ce groupe font l’objet de mesure de quarantaine très sévère lors de leur introduction et considérant qu’elle n’a jamais été trouvée ou signalée sur le continent Africain auparavant, une enquête couvrant près de 3000 km à travers les principales localités productrices de fruits du Kenya a immédiatement été lancée afin de déterminer l’aire d’extension de cette nouvelle espèce invasive. Nous présentons des résultats sur la découverte de la mouche ainsi que sur l’enquête, et discutons des effets potentiels de cette mouche sur l’industrie horticole de l’Afrique de l’Est.

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.

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.

Parasitoids of medfly, Ceratitis capitata, and related tephritids in Kenyan coffee: a predominantly koinobiont assemblage

Arabica coffee was sampled from two sites in the central highlands of Kenya (Rurima, Ruiru) and one site on the western side of the Rift Valley (Koru). Three species of ceratitidine Tephritidae, Ceratitis capitata (Wiedemann), C. rosa Karsch and Trirhithrum coffeae Bezzi, were reared from sites in the central highlands, and an additional species, C. anonae Graham, was recovered from the western-most site. Ten species of parasitic Hymenoptera were reared from these tephritids. The parasitoid assemblage was dominated by koinobionts. Eight of the species are koinobiont endoparasitoids, but only one idiobiont larval ectoparasitoid was reared, and only one idiobiont pupal endoparasitoid. The effects of sampling bias on determination of parasitoid assemblage size associated with concealed hosts are discussed. The potential for use of these parasitoids in biological control is also discussed. Most of the parasitoid species recovered during this study are capable of developing on C. capitata, while several also attack C. rosa. Both flies are notorious pests of tropical and subtropical fruits.

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.

Geographic Distribution, Host Fruit, and Parasitoids of African Fruit Fly Pests Ceratitis anonae, Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis rosa (Diptera: Tephritidae) in Kenya

Abstract The fruit of indigenous, cultivated, and naturalized exotic plants was sampled in Kenya to determine the geographical and host plant distributions of the fruit fly pests Ceratitis anonae (Graham), Ceratitis cosyra (Walker), Ceratitis fasciventris (Bezzi), and Ceratitis rosa Karsch. In 1997, cultivated and wild fruit were sampled on the Kenya coast to determine seasonal patterns of host use by C. cosyra and C. rosa. From 1999 to 2004, the sampling effort was expanded to forested areas of the central and western highlands and to all fruit-infesting tephritids. Together, the four pest tephritid species were reared from 5.1% of 3,794 fruit collections, the latter making up 116 families and 882 species of host plants. C. anonae, C. cosyra, C. fasciventris, and C. rosa were reared throughout Kenya, from 14, 9, 30, and 28 plant species, respectively. Fifty-two of these plants represented previously unknown hosts for one or more of the tephritids. C. anonae was restricted to habitats west of the Gregory Rift Valley. C. fasciventris was found in western and central Kenya, but it was not reared from coastal fruit collections. C. rosa occurred at the coast and in central highland fruit. We provide evidence for the recent introduction of C. rosa into the central highlands. Only C. cosyra was found in habitats located in all three of the major regions sampled. Although distributed over a wider geographic area than the other species, C. cosyra had a markedly restricted host range. On the coast, C. cosyra used wild fruit, primarily Sclerocarya birrea (A. Rich.) Hochst. (Anacardiaceae), as hosts when mango was not fruiting. C. rosa was absent from our mango samples but attacked common guava and wild fruit species. Overall, C. anonae, C. fasciventris, and C. rosa had similar host profiles, but each also had unique hosts in which infestations were usually heavy. Members of the Sapotaceae and Annonaceae were the most important wild hosts of these three species. Within these families, host fruit were partitioned among these tephritid species and the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). The availability of wild host fruit, sometimes supplemented by naturalized invasive plants or cultivated fruit, provided for year-round breeding of C. anonae and C. fasciventris, whereas indigenous fruit were sufficient for breeding of C. rosa and C. cosyra during most of the year. Opiine braconids were reared from all four species of Ceratitis.

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.

Comparative studies on the fecundity, egg survival, larval feeding, and development of Chilo partellus and Chilo orichalcociliellus (Lepidoptera: Crambidae) on five grasses.

Abstract Chilo partellus (Swinhoe) and Chilo orichalcociliellus Strand (Lepidoptera: Crambidae) are gramineous stem borers that occur sympatrically in the southern coastal area of Kenya. Evidence over a 30-yr period indicates that the indigenous stem borer, C. orichalcociliellus, is being gradually displaced by the exotic stem borer, C. partellus. Comparative laboratory studies were conducted in several large-stemmed grasses to examine factors that may be involved in the displacement of C. orichalcociliellus, and to examine other possible effects of the invasion of C. partellus into Kenya. C. partellus had a higher fecundity than C. orichalcociliellus at 25 and 28°C, but not at 31°C. In addition, more C. partellus than C. orichalcociliellus eggs survived to the first instar. C. partellus larvae developed faster than C. orichalcociliellus in maize and sorghum. In this shorter developmental time, C. partellus consumed more maize than C. orichalcociliellus, but both species consumed equal amounts of sorghum. On a daily basis, C. partellus consumed more maize and sorghum than C. orichalcociliellus. A few C. orichalcociliellus survived to the pupal stage in napier and guinea grasses, whereas no C. partellus survived. The shorter developmental period of C. partellus may give this species a competitive advantage over the slower developing C. orichalcociliellus. However, the ability of C. orichalcociliellus to complete development in two native grasses in which C. partellus did not survive may provide a refuge that has allowed C. orichalcociliellus to escape extirpation from the coastal area of Kenya.

Population genetics of the potentially invasive African fruit fly species, Ceratitis rosa and Ceratitis fasciventris (Diptera: Tephritidae)

A set of 10 microsatellite markers was used to survey the levels of genetic variability and to analyse the genetic aspects of the population dynamics of two potentially invasive pest fruit fly species, Ceratitis rosa and C. fasciventris, in Africa. The loci were derived from the closely related species, C. capitata. The degree of microsatellite polymorphism in C. rosa and C. fasciventris was extensive and comparable to that of C. capitata. In C. rosa, the evolution of microsatellite polymorphism in its distribution area reflects the colonization history of this species. The mainland populations are more polymorphic than the island populations. Low levels of differentiation were found within the Africa mainland area, while greater levels of differentiation affect the islands. Ceratitis fasciventris is a central–east African species. The microsatellite data over the Uganda/Kenya spatial scale suggest a recent expansion and possibly continuing gene flow within this area. The microsatellite variability data from C. rosa and C. fasciventris, together with those of C. capitata, support the hypothesis of an east African origin of the Ceratitis spp.