M. De Meyer

Seasonality and host utilization of the invasive fruit fly, Bactrocera invadens (Dipt., Tephritidae) in central Tanzania

Abstract:  The temporal occurrence of the invasive and economically important pest fruit fly, Bactrocera invadens was studied in three agro‐ecological areas of Morogoro Region, central Tanzania, during 2004–2005. Weekly and monthly trappings were carried out with methyl eugenol, protein bait and synthetic food attractant. Bactrocera invadens was permanently present at low and mid‐altitudes (380–520 m a.s.l.) with peak periods coinciding with the fruiting season of mango (Mangifera indica) and guava (Psidium guajava). At high altitude (1650 m a.s.l.) its incidence was only temporal and apparently the result of dispersal from lower altitudes after the mango fruiting season. Rearing results showed mango, loquat (Eriobotrya japonica), guava and grapefruit (Citrus × paradisi) to be the favoured commercial host fruits. Other Citrus species, cucurbits, papaya (Carica papaya) and avocado (Persea americana) were less favoured.

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.

Host range and distribution of fruit-infesting pestiferous fruit flies (Diptera, Tephritidae) in selected areas of Central Tanzania

The host range of major fruit fly pests in Central Tanzania was evaluated from October 2004 to October 2006. Samples of 48 potential hosts were collected and incubated for fruit fly emergence. Bactrocera invadens was the dominant species in incidence expressed as the ratio of infested to total number samples collected, as well as infestation rate, expressed as number of flies emerging per unit weight. Eight new host fruits are reported. Infestation by native pests, such as Ceratitis capitata and C. cosyra, was minor compared to B. invadens. Ceratitis rosa was the dominant species in temperate fruits, and Cucurbitaceae were mainly infested by Bactrocera cucurbitae, a specialized cucurbit feeder. Among commercial fruits, high infestation incidences were observed in mango and guava, but they decreased throughout the fruiting season. Low infestation rates were observed in all Citrus species and in avocado, indicating these fruits as poor hosts for the studied fruit fly pests in this region. Widespread availability and abundance of fruit species studied here ensures year-round breeding of B. invadens. Seasonal infestation differs, with mango being the most important host in October to January, while guava being important from February to August. Tropical almond showed very high incidence and infestation rate for B. invadens and might act as an important reservoir host, bridging the fruiting seasons of mango and guava. Soursop acts as an important host for C. cosyra after the mango season. Ceratitis capitata is a pest of minor importance of the commercial fruits studied in this region.

Recovering full DNA barcodes from natural history collections of Tephritid fruitflies (Tephritidae, Diptera) using mini barcodes

The family of Tephritid fruit flies (Tephritidae, Diptera) is composed of more than 4000 species and more than 350 are of economic importance (EI). The Tephritid Barcoding Initiative (TBI) aims at obtaining DNA barcodes for all EI species and the majority of their congeners. Dry pinned specimens from natural history collections are an important resource for reference material, but were often collected decades ago. We observed a strong decrease in the success rate of obtaining a full COX1 DNA barcode (658 bp), with an increasing age of the specimens. Obtaining full barcodes is often not possible using standard protocols. We developed a universal Tephritid primer set for multiple overlapping mini‐barcodes that allows reconstructing the full COX1 DNA barcode. These newly developed primers and the corresponding protocol will facilitate the utilization of the extensive natural history collection by the TBI consortium.

Macrogeographic population structuring in the cosmopolitan agricultural pest Bactrocera cucurbitae (Diptera: Tephritidae).

The macrogeographic population structure of the agricultural pest Bactrocera cucurbitae (Diptera: Tephritidae) was investigated in order to identify the geographic origin of the species and reconstruct its range expansion. Individuals of B. cucurbitae were collected from 25 worldwide‐distributed localities (n = 570) and genotyped at 13 microsatellite loci. The Bayesian clustering reveals that B. cucurbitae can be subdivided into five main groups corresponding to populations from (i) the African continent, (ii) La Réunion, (iii) Central Asia, (iv) East Asia and (v) Hawaii. The proportions of inter‐regional assignments and the higher values of genetic diversity in populations from Pakistan, India and Bangladesh suggest that B. cucurbitae originated in Central Asia and expanded its range to East Asia and Hawaii on one hand and to Africa and the islands of the Indian Ocean on the other. A number of outliers (10–19 specimens according to different clustering algorithms) show high levels of admixture (Q > 0.70) with populations from different regions and reveal complex patterns of inter‐regional gene flow. Anthropogenic transport is the most plausible promoter of this large‐scale dispersal. The introduction of individuals from geographically distant sources did not have a relevant role in the most recent African invasions, which originated from the expansion of local populations. These results could provide a useful background to better evaluate invasion risks and establish priorities for the management of this cosmopolitan agricultural pest.

Uncovering the tracks of a recent and rapid invasion: the case of the fruit fly pest Bactrocera invadens (Diptera: Tephritidae) in Africa.

Phytophagous insects of the genus Bactrocera are among the most economically important invasive fruit fly pests. In 2003, an unknown Bactrocera species was found in Kenya. First identified as an ‘aberrant form’ of the Asian B. dorsalis complex, it was later recognized as a new species, Bactrocera invadens. Within 2 years of its discovery, the species was recorded in several African countries, becoming an important quarantine pest. As this invasive fly was discovered only recently, no data are available on its invasion pattern in Africa. This pilot study attempts to infer from genetic data the dynamic aspects of the African invasion of this pest. Using microsatellite markers, we evaluated the level of genetic diversity and the extent of common ancestry among several African populations collected across the invaded areas. A sample from the Asian Sri Lankan population was analysed to confirm the Asian origin of this pest. Genetic data cast no doubt that Sri Lanka belongs to the native range, but only a small percentage of its genotypes can be found in Africa. African populations display relatively high levels of genetic diversity associated with limited geographical structure and no genetic footprints of bottlenecks. These features are indicative of processes of rapid population growth and expansion with possible multiple introductions. In the span of relatively few years, the African invasion registered the presence of at least two uncorrelated outbreaks, both starting from the East. The results of the analyses support that invasion started in East Africa, where B. invadens was initially isolated.

Molecular evaluation of nominal species in the Ceratitis fasciventris, C. anonae, C. rosa complex (Diptera: Tephritidae)

Ceratitis fasciventris, C. anonae and C. rosa form a complex of economically important fruit fly pests infesting a variety of crops in African countries. Hitherto only adult males of these species can be distinguished easily by morphological characters. Other stages cannot, and for some taxa the taxonomic interpretation and species boundaries remain unclear. In order to clarify phylogenetic relationships and taxonomic status of these species, sequences of mitochondrial (16S, COI, ND6) and nuclear markers (period, ITS1) were analysed in specimens of the three morphospecies throughout the distribution of the complex. Maximum likelihood trees did not recover monophyletic groups corresponding to the morphospecies. Conversely, ND6 and COI divided West African C. fasciventris specimens in two consistent and bootstrap supported clades, involving specimens from Benin and from Mali/Ivory Coast, while the nuclear gene fragments per and ITS1 recovered a well-supported clade corresponding to C. fasciventris from Kenya/Uganda. Hence, the phylogenetic relationships and taxonomic interpretation of the complex appear more intricate than previously hypothesised. The current molecular data do not allow to identify C. fasciventris, C. anonae and C. rosa as distinct phylogenetic species but rather suggest that the morphospecies C. fasciventris is itself a complex of cryptic taxa.

Cryptic diversity and gene flow among three African agricultural pests: Ceratitis rosa, Ceratitis fasciventris and Ceratitis anonae (Diptera, Tephritidae).

The ‘Ceratitis FAR complex’ is a species complex of African fruit flies (Diptera, Tephritidae) including the major agricultural pest Ceratitis rosa and the morphologically similar Ceratitis fasciventris and Ceratitis anonae. To resolve their intra‐ and interspecific genetic relationships and to estimate gene flow within this complex, we surveyed allelic variation at 16 microsatellite loci in 27 African populations of the three morphospecies. Interpopulation genetic distances and individual Bayesian assignments distinguished five genotypic clusters: two involving C. rosa (R1, R2; that may occur in sympatry), two involving C. fasciventris (F1, F2; with parapatric distributions) and one involving C. anonae (A). Intra‐ and interspecific patterns of genetic differentiation were not hierarchically structured and genetic differentiation between conspecific clusters (F1–F2 and R1–R2) was higher or comparable with differentiation between heterospecific clusters (e.g. F1‐A or R2‐A). In some cases, gene flow estimates among morphospecies or among heterospecific genotypic clusters were significantly different from zero, showing the lack of reproductive isolation. Genetic differentiation between genotypic clusters was partly supported by morphological differences observed a posteriori in male secondary sexual characters. These results suggest important revisions to current models of ecological niche requirements and invasion risk of the major agricultural pest C. rosa and provide a basis for a taxonomic re‐interpretation of the FAR complex.

Detection of the solanum fruit fly, Bactrocera latifrons (Hendel) in Tanzania (Dipt., Tephritidae).

Abstract:  The presence of the Solanum fruit fly, Bactrocera latifrons, in Africa is reported for the first time, based on trapped and reared specimens in Tanzania. Two new host records, Solanum aethiopicum and Solanum macrocarpon, are reported.

Molecular Identification of Ceratitis capitata (Diptera: Tephritidae) using DNA Sequences of the COI Barcode Region

ABSTRACT The utility of the cytochrome oxidase I gene barcode region for diagnosis of the Mediterranean fruit fly, Ceratitis capitata (Weidemann), is evaluated using African fruit fly collections. The method fails to discern C. capitata from its close relative Ceratitis caetrata Munro, based on genetic distances, parsimony networks, or nucleotide diagnostic characters observed in the DNA barcode sequences. When treated as a single taxon, it is possible to discern the C. capitata + C. caetrata lineage from other Ceratitis species. Levels of intraspecific diversity vary within the genus Ceratitis and multiple copies of the mitochondrial gene are reported for Ceratitis cosyra (Walker). The DNA barcoding method based on genetic distance is compared with a molecular identification method using restriction fragment length polymorphism. The DNA barcode and restriction fragment-length polymorphism methods provide similar identification results, but the DNA sequence information is more suitable for quantitative analysis of the information.