The Effects of Selection for early (day) and late (dusk) Mating lines of Hybrids of Bactrocera tryoni and Bactrocera neohumeralis

Bactrocera neohumeralis and Bactrocera tryoni are closely related tephritid fruit fly species. B. neohumeralis mates throughout the day (in bright light) and B. tryoni mates at dusk. The two species can also be distinguished by the colour of their calli (prothoracic sclerites) which are brown and yellow, respectively. The F1 hybrids can mate both in bright light just before dusk and during dusk and have calli that are partly brown and partly yellow. The F2 hybrids have a wider range of callus patterns and mating occurs more widely in the day as well as at dusk. We directly selected hybrid stocks for mating time, creating 'early' (day-mating) and 'late' (dusk-mating) lines. As an apparently inadvertent consequence, the two types of line respectively had predominantly brown and predominantly yellow calli and thus came to closely resemble the original two species in both behaviour and appearance. Lines that were evenly selected (half for day and half for dusk) essentially retained the mating pattern of F2 hybrids. Selection for callus colour alone also affected the distribution of mating times in a predictable way. We propose a genetical model to account for the results and discuss them in the light of the apparent maintenance of species integrity in nature.     

The likely fate of hybrids of Bactrocera tryoni and Bactrocera neohumeralis

Bactrocera tryoni (Froggatt) and B. neohumeralis (Hardy) (Diptera: Tephritidae) are sympatric species which hybridise readily in the laboratory yet remain distinct in the field. B. tryoni mates only at dusk and B. neohumeralis mates only during the day, but hybrids can mate at both times. We investigated the inheritance of mating time in successively backcrossed hybrid stocks to establish whether mating with either species is more likely. The progeny of all backcrosses to B. tryoni mated only at dusk. The majority of the progeny of the first and a minority of the progeny of the second backcross to B. neohumeralis also mated at dusk, but the third successive B. neohumeralis backcross produced flies that mated only during the day. This trend towards dominance of the B. tryoni trait was also reflected in a diagnostic morphological character. We discuss the possible genetic background for these phenomena and propose that unidirectional gene flow might explain how the two species remain distinct in the face of natural hybridisation.     

Genetic delineation of sibling species of the pest fruit fly Bactocera (Diptera: Tephritidae) using microsatellites

Using a large set of microsatellites, the genetic relationships between three closely related Australian fruit fly species, Bactrocera tryoni (Froggatt), B. neohumeralis (Hardy) and B. aquilonis(May) were investigated. Bactrocera tryoni and B. neohumeralis are sympatric, while B. aquilonisis allopatric to both. The sympatric species, B. tryoni and B. neohumeralis, were found to be genetically distinct. It is likely that despite differences in mating time between these two species, some gene flow still occurs. In contrast, the sibling species B. tryoni and B. aquilonis were found to be closely related, despite allopatry. The level of genetic divergence was similar to that found within eastern Australian populations of B. tryoni. Consideration of all available genetic data suggests that this similarity is not due to recent (i.e. within the last 30 years) displacement of B. aquilonis by B. tryoni from the B. aquilonis region (north-western Australia). Instead the data suggests that, at least in the areas sampled, asymmetrical hybridization may have occurred over a longer timescale.     

Attributes pertinent to overwintering potential do not explain why Bactrocera neohumeralis (Hardy) (Diptera: Tephritidae) does not spread further south within the geographical range of B. tryoni (Froggatt)

Abstract  The geographical range of Bactrocera neohumeralis does not extend as far south as that of its sibling species, B. tryoni . However, there was no evidence of any difference between the two species in terms of physiological limitation to southerly spread when comparisons were made of low temperature torpor thresholds of adults, survival time of adults at −4°C and development rates of all stages in either warm or cool regimes. The survival schedule of the two species was similar in the laboratory and also in the moderately cold conditions experienced by caged cohorts that were exposed to winter field temperatures between late April and early November at Richmond, New South Wales (500 km south of the usual southerly limit of B. neohumeralis ). Overwintered cohorts of both species laid similar numbers of eggs in September in terms of eggs per emerged female (an indicator of the reproductive potential). However, because the proportion of B. tryoni surviving to the period of 1–15 September was less than half that for B. neohumeralis , the production per surviving female was more than double in B. tryoni . The possibility of the southerly spread of B. neohumeralis being limited by an Allee effect is discussed.     

Close genetic similarity between two sympatric species of tephritid fruit fly reproductively isolated by mating time

Abstract.— Two sibling species of tephritid fruit fly, Bactrocera tryoni and B. neohumeralis , occur sympatrically throughout the range of B. neohumeralis in Australia. Isolation between the two species appears to be maintained by a difference in mating time: B. tryoni mates at dusk, whereas B. neohumeralis mates during the middle of the day. A morphological difference in humeral callus color also distinguishes the two species. Despite clear phenotypic evidence that B. tryoni and B. neohumeralis are distinct species, genetic differentiation as measured by four markers–nuclear DNA sequences from the white gene and the ribosomal internal transcribed spacer (ITS2), and mitochondrial DNA sequences from the cytochrome b (cytb) and cytochrome oxidase subunit II (COII) genes–is very small. Minor fixed differences occur in the ITS2 sequence, however, in all other cases the two species exhibit a high level of shared polymorphic variation. The close genetic similarity suggests either that speciation has occurred very rapidly and recently in the absence of any mitochondrial DNA sorting or that the sharing of polymorphisms is due to hybridization or introgression. A third species within the tryoni complex, B. aquilonis , is geographically isolated. Bactrocera aquilonis is also genetically very similar, but in this case there is clear differentiation for the mitochondrial loci. The three species form a group of considerable interest for investigation of speciation mechanisms.     

Australian Distribution of 17 Species of Fruit Flies (Diptera: Tephritidae) Caught in Cue Lure Traps in February 1994

We report the first widespread survey of tephritid fruit flies attempted in a single time period. 1,471 cue lure traps caught 17 species, and extensions to previously recorded geographical ranges were detected for seven of them: Bactrocera tryoni, B. neohumeralis, B. frauenfeldi, B. aeroginosa, Dacus absonifascies, D. aequalis and D. newmani. the traps also unexpectedly caught several B. cacuminata and also both males and females of Dirioxa pornia and Ceratitis capitata. the geographical variation in the relative abundance of B. tryoni and B. neohumeralis in the region of their co-occurrence was in substantial agreement with earlier estimates. the regional variation in abundance of B. tryoni in the eastern states was in accordance with the predictions of a published bioclimatic model. Furthermore, the spread of this species (expected from the model) to several locations in the Northern Territory is recorded here for the first time.     

The influence of soil type and moisture on pupal survival of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  Larvae of the Queensland fruit fly, Bactrocera tryoni , pupate in the soil, but the influence of soil variables on B. tryoni pupal mortality is not known. For other tropical tephritid species, soil moisture has been identified as a major pupal mortality factor. In the laboratory, we tested the effects of soil moisture and soil type on pupal survival through a factorial experiment which used three soil types (loamy sand, loam, sandy clay) and seven soil moisture levels (0%, 10%, 25%, 50%, 75%, 90% and 100%). Minor, but significant, differences in pupal mortality were observed between the soil types, but the most significant factor affecting pupae was extremes of soil moisture. Eighty-five percent pupal mortality occurred at 0% soil moisture and 30% mortality at 100% soil moisture: very low levels of mortality occurred at all intermediate levels. We detected a significant interaction between soil type and moisture level but cannot explain it. In a follow-up experiment, we demonstrated that prepupal wandering larvae of B. tryoni could discriminate between different moisture levels, with significantly greater pupation in loam soil at 75% soil moisture than at either 0% or 100% soil moisture. Results are used to modify a pupal mortality/soil moisture equation used in a recently published DYMEX model of B. tryoni population dynamics .     

Enzyme polymorphism and species discrimination in fruit flies of the genus Dacus (Tephritidae).

Sympatric populations of D. tryoni and D. neohumeralis are difficult to completely distinguish taxonomically. Using five pigmentation characters, each of some taxonomic value, a small proportion of individuals cannot be assigned to either species nor definitely classified as hybrids. To aid in species discrimination and hybrid identification gene frequencies in natural populations were estimated at three polymorphic protein loci, an alcohol dehydrogenase (Adh), an octanol dehydrogenase (Odh) and an esterase (E-2). Samples of flies were taken from four sites spread over 1200 miles along the Australian eastern coast. Within each species allelic frequencies at each locus were largely the same at all localities. Consistent differences in gene frequencies between species occurred at all three loci, strongly supporting the hypothesis of two distinct gene pools. The Adh locus best discriminated between species with a unique allele occurring in D. neohumeralis at a frequency of 0-85. None of the loci showed complete differentiation and hence it was not possible to find a quick and easy method to distinguish the species nor to detect field hybrids. Directional selection of laboratory populations for a change in callus colour (the best pigmentation character for separating the species) indicated that at the Adh and E-2 loci frequencies of major alleles were not genetically associated with major genes for callus colour. Thus genotype determination at these loci when considered together with pigmentation characters may be valuable taxonomically for further distinguishing between the species.     

Short-range dispersal of recently emerged males and females of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) monitored by sticky sphere traps baited with protein and Lynfield traps baited with cue-lure

Abstract  Dispersal of immature male and female Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), was assessed over a period of 1 week from a single release point on three separate occasions using an array of Lynfield traps baited with cue-lure and odouriferous yellow or black sticky spheres baited with food lure (protein autolysate). Lynfield traps recaptured males; yellow or black spheres recaptured both sexes in approximately equal proportions, although at a much lower rate. As a percentage of the recapture rate for males by Lynfield traps, the mean recapture rate for yellow spheres ranged from 1.0% to 7.5% for males and 0.7% to 4.0% for females, whereas the recapture rates for black spheres ranged from 0.4% to 3.6% and 0.6% to 1.8%, respectively. The rate of recapture of sterile male flies was greater than that of unsterilised flies; this may have been due to a faster maturation rate in sterile males or because a greater proportion of them remained within the trap array rather than dispersing. There was no significant trend in recapture rate with distance from the release point to the edge of the array (88 m), except in the case of females on sticky traps where no trend was detected between 19 and 88 m. These results lend support to assumptions made about the distribution of males and females with respect to the minimum breeding density of fruit fly propagules invading a fly-free zone, and the method chosen to distribute sterile B. tryoni for the sterile insect technique.     

Influence of male aggregation size on female visitation in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The importance of male aggregation size for female visitation and initiation of male pheromone-calling was investigated in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) using artificial male aggregations in large laboratory cages. Female B. tryoni visited the largest aggregation more frequently than single males in association with a higher proportion of calling males, but there was no correlation between aggregation size and female visitation. Female B. tryoni had a limited capacity to perceive a difference between the number of calling males. Calling propensity of male B. tryoni was increased by the presence of conspecific males. Increased calling propensity in larger groups of male B. tryoni may be due to social facilitation of male calling behaviour. Female visitation at aggregations was only weakly associated with male calling, suggesting that aggregation size and the number of pheromone-calling males are not the only factors important in locating mates in B. tryoni , and it is possible that low-density populations could persist so long as females can encounter single males.     

REDUCTION IN TRAP CAPTURES OF FEMALE FRUIT FLIES (DIPTERA: TEPHRITIDAE) WHEN SYNTHETIC MALE LURES ARE ADDED

Abstract
Captures of females of 4 species of fruit flies on or in both coloured sticky traps and protein baited traps were reduced by synthetic male attractants. Captures of female Dacus tryoni (Froggatt), Dacus neohumeralis Hardy and Dacus cacuminatus (Hering) were reduced by both cuelure and methyl eugenol, and female Ceratitis capitata (Weidemann) by Capilure®. These effects were sufficient to overcome the normally attractive stimuli of colour and odours associated with yeast.     

THESIS ABSTRACT

Geographical distribution, seasonal abundance and habitat preference of fruit fly species (Diptera: Tephritidae) in South-East Queensland, with special reference to Bactrocera tryoni (Froggatt) and Bactrocera neohumeralis (Hardy) . S. Raghu.     

A new family of Strepsiptera parasitizing fruit flies (Tephritidae) in Australia

ABSTRACT. A new family of Strepsiptera (Dipterophagidae) from Australia containing a new genus and species ( Dipterophagus daci ) is described and figured. This is the first species of Strepsiptera to be described as a parasite of Diptera. D.daci is recorded from ten fruit fly species, viz Dacus aquilonis (May), D.bellulus Drew, D.cacuminatus (Hering), D.decurtans (May), D.mayi Hardy, D.neohumeralis Hardy, D.peninsularis Drew, D.tenuifascia (May), D.tryoni (Froggatt) and Dacus sp. n. ( near musae ). Gregarious parasitism (two to six adult parasites per fly) is common.     

Influence of fruit traits on oviposition preference and offspring performance of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) on three tomato (Lycopersicon lycopersicum) cultivars

Abstract  In Queensland, three tomato ( Lycopersicon lycopersicum ) cultivars, Grosse Lisse, Roma and Cherry, are infested by Queensland fruit fly, Bactrocera tryoni (Froggatt). In this study, we examined if there was a correlation between oviposition preference and offspring performance of B. tryoni among the three tomato cultivars. We also investigated host plant traits that may explain any variation in preference and performance. Choice and no-choice experiments were carried out under laboratory conditions. A positive correlation between oviposition preference and offspring performance of B. tryoni was observed in the three tomato cultivars. Grosse Lisse and Roma cultivars were highly preferred by B. tryoni over Cherry cultivar. Performance (measured as proportion of eggs developing to the pupal stage) was significantly higher in Grosse Lisse and Roma cultivars than in Cherry cultivar. The pericarp toughness of Cherry cultivar appears responsible for its low rate of infestation, while the presence of 2-butanol and 1,4-butanediamine in Roma and Grosse Lisse, respectively, may partly be responsible for the high oviposition preference shown by B. tryoni towards these cultivars.     

Impact of introduction of Bactrocera dorsalis (Diptera: Tephritidae) and classical biological control releases of Fopius arisanus (Hymenoptera: Braconidae) on economically important fruit flies in French Polynesia

Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), was discovered on Tahiti Island in July 1996. Eradication programs were conducted from 1997 to 2001, but failed. From 1998 to 2006, B. dorsalis was recovered from 29 different host fruit from the five Society Islands: Tahiti, Moorea, Raiatea, Tahaa, and Huahine. Analysis of coinfestation patterns by B. dorsalis, Bactrocera tryoni (Froggatt), and Bactrocera kirki (Froggatt) suggested B. dorsalis had displaced these two species and become the most abundant fruit fly in coastal areas. To suppress B. dorsalis populations, a classical biological control program was initiated to introduce the natural enemy Fopius arisanus (Sonan) (Hymenoptera: Braconidae) into French Polynesia from Hawaii. Wasps were released and established on Tahiti, Moorea, Raiatea, Tahaa, and Huahine Islands. In guava, Psidium guajava L., collections for Tahiti, F. arisanus parasitism of fruit flies was 2.1, 31.8, 37.5, and 51.9% for fruit collected for 2003, 2004, 2005 and 2006, respectively. Based on guava collections in 2002 (before releases) and 2006 (after releases), there was a subsequent decrease in numbers of B. dorsalis, B. tryoni, and B. kirki fruit flies emerging (per kilogram of fruit) by 75.6, 79.3, and 97.9%, respectively. These increases in F. arisanus parasitism and decreases in infestation were similar for other host fruit. Establishment of F. arisanus is the most successful example of classical biological control of fruit flies in the Pacific area outside of Hawaii and serves as a model for introduction into South America, Africa, and China where species of the B. dorsalis complex are established.     

Attraction of fruit flies of the genus Bactrocera to colored mimics of host fruit

In tests on feral populations of polyphagous Bactrocera tryoni (Froggatt) adults on host guava trees, both sexes were significantly more attracted to Tangletrap‐coated 50 mm diameter spheres colored blue or white than to similar spheres colored red, orange, yellow, green, or black or to Tangletrap‐coated 50 mm diameter yellow‐green guava fruit. In contrast, in tests on feral populations of oligophagous Bactrocera cacuminata (Hering) on host wild tobacco plants, both sexes were significantly more attracted to Tangletrap‐coated 15 mm diameter spheres colored orange or yellow than to other colors of spheres or to Tangletrap‐coated 15 mm diameter green wild tobacco fruit. Both sexes of both tephritid species were significantly more attracted to blue (in the case of B. tryoni) or orange (in the case of B. cacuminata) 50 mm spheres displayed singly than to blue or orange 15 mm spheres displayed in clusters, even though fruit of wild tobacco plants are borne in clusters. Finally, B. tryoni adults were significantly less attracted to non‐ultraviolet reflecting bluish fruit‐mimicking spheres than to bluish fruit‐mimicking spheres having a slightly enhanced level of ultraviolet reflectance, similar to the reflectance of possible native host fruit of B. tryoni, whose bluish skin color is overlayed with ultraviolet‐reflecting waxy bloom. Responses to fruit visual stimuli found here are discussed relative to responses found in other tephritid species.     

Hybrid queen butterflies from the cross Danaus chrysippus (L.) × D. gilippus (Cramer):confirmation of species status for the parents and further support for Haldane's Rule

A cross between queen butterflies of the Palaeotropical species Danaus chrysippus and the Neotropical D. gilippus was achieved with difficulty in both directions. Only one progeny ( N  = 70) was reared comprising sterile males and inviable females in a precisely 1 : 1 ratio. Both prezygotic and postzygotic barriers to gene flow are strong. The result supports Haldane's Rule, to which we propose a minor amendment. The F₁ hybrids were intermediate for background colour between the brown (genotype BB ) of gilippus and orange (genotype bb ) of chrysippus . Most F₁ pattern characters were also intermediate. In polymorphic chrysippus populations, because Bb heterozygotes are brown, or nearly so, we suggest the B allele may have evolved towards dominance in sympatry. Hybrid males show positive heterosis for body size. The close similarity of male genitalia between the allopatric, genetically distant species chrysippus  and gilippus , compared to their divergence between gilippus and its largely sympatric sister species eresimus , suggest that reinforcement of sexual isolation or reproductive character displacement have evolved in sympatry.  © 2002 The Linnean Societyof London, Biological Journal of the Linnean Society , 2002, 76 , 535–544.     

Genetic consequences of domestication and mass rearing of pest fruit fly Bactrocera tryoni (Diptera: Tephritidae)

Tephritid fruit flies, an important pest of horticulture worldwide, are increasingly targeted for control or eradication by large-scale releases of sterile flies of the same species. For each species treated, strains must be domesticated for mass rearing to provide sufficiently large numbers of individuals for releases. Increases in productivity of domesticated tephritid strains are well documented, but there have been few systematic studies of the genetic consequences of domestication in tephritids. Here, we used nine DNA microsatellite markers to monitor changes in genetic diversity during the early generations of domestication in replicated lines of the fruit fly Bactrocera tryoni (Froggatt) (Diptera: Tephritidae). The observed changes in heterozygosity and allelic richness were compared with the expected changes in heterozygosity generated by a stochastic simulation including genetic drift but not selection. The results showed that repeatable genetic bottlenecks occur in the early generations and that selection occurs in the later generations. Furthermore, using the same simulation, we show that there is inadvertent selection for increased productivity for the entire life on a mass-rearing colony, in addition to intentional selection for increased productivity. That additional selection results from the common practice of establishing the next generation of the breeding colony from a small proportion of one day's pupae collection (the pupal raffle). That selection occurs during all generations and acts only on fecundity variation. Practical methods to counter that unavoidable loss of genetic diversity during the domestication process in B. tryoni are discussed.     

Genetic and molecular markers of the Queensland fruit fly, Bactrocera tryoni

Twenty-six microsatellite markers, along with two restriction fragment length polymorphism (RFLP) markers and three morphological markers, have been mapped to five linkage groups, corresponding to the five autosomes of the Queensland fruit fly, Bactrocera tryoni. All these molecular and genetic markers were genotyped in three-generation pedigrees. Eight molecular markers were also localized to the salivary gland polytene chromosomes by in situ hybridization. This provides a substantial starting point for an integrated genetic and physical map of B. tryoni.