DNA microsatellite analysis of naturally occurring colour intermediates between Bactrocera tryoni (Froggatt) and Bactrocera neohumeralis (Hardy) (Diptera: Tephritidae)

Abstract  The sympatric tephritid fruit flies Bactrocera tryoni (Froggatt) (Queensland fruit fly) and B. neohumeralis (Hardy) differ in time of mating and for the colour of the humeral callus ('shoulder pad'), which is typically entirely yellow in B. tryoni and typically entirely brown in B. neohumeralis . Field collections in sympatric regions usually include at least 1% of individuals whose humeral calli show mixed patches of yellow and brown ('intermediates'). Over 40 years, a number of studies have debated the possibility that these intermediates are interspecific hybrids. In the present study, we have used microsatellites to show that few if any of these intermediates are hybrids. Instead, most variation humeral callus appears to be confined to one species, B. tryoni . We discuss these results in the context of geneflow between the two species and suggest directions for future research.     

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.     

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.     

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.     

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.     

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.     

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 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 .     

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.     

Life history parameters of Orius sauteri (Poppius) (Het., Anthocoridae) reared on Ephestia kuehniella eggs and the minimum amount of the diet for rearing individuals

Abstract: Mass rearing techniques are essential elements in developing a reliable and economic biological control method for Orius sauteri, a promising indigenous Orius species for biological control of Thrips palmi in greenhouses. Orius sauteri was reared on Ephestia kuehniella eggs and life history parameters and the minimum number of E. kuehniella eggs for rearing O. sauteri individuals were measured at 25°C and L16:D8. Developmental time during the nymphal stages was about 14 days for both sexes. Total fecundity and female longevity were 103.9 and 27.9 days, respectively. The intrinsic rate of natural increase per day was 0.115. An individual requires at least 30 eggs for nymphal development and survival and 20 eggs for adult oviposition and survival per four days.     

Population dynamics and reproductive aspects of Porcellio albinus (Isopoda,Oniscidea) of Zarat (Gabes,Tunisia)

Porcellio albinus is a nocturnal species inhabiting xeric habitats in Tunisia. Size distribution, population dynamics, breeding activity, fecundity, fertility, reproductive allocation (RA), and life span were studied to establish the reproductive strategy of a population of this species in the sandy coastal area of Zarat. Monthly sampling was undertaken between July 2012 and November 2013. The overall sex-ratio of P. albinus was female biased. The relative abundance of adults, much more representative than that of juveniles, might be the consequence of the absence of specimens less than 8 mm in size in all samples collected outside the burrows at night. Reproduction is seasonal in P. albinus with two breeding seasons, the first one from March to June and the second in September. Five cohorts were recruited per year, four during the spring and one during the fall. The number of eggs per female ranged from 13 to 67 and was positively correlated with female body size. Fertility varied from 5 to 39 mancae. Compared to other xeric species, RA was low to produce a brood, but high to release a single manca. The results provide a better understanding of the reproductive behavior of this terrestrial isopod, living under severe environmental conditions.     

Host preference and host suitability in an egg-pupal fruit fly parasitoid, Fopius arisanus (Sonan) (Hym., Braconidae)

The relative oviposition rate of the parasitoid Fopius arisanus (Sonan) was investigated across three frugivorous tephritid species, Bactrocera tryoni Froggart, Bactrocera jarvisi (Tryon) and Bactrocera cucumis French. Choice and no-choice tests were both used. The suitability of these three species for sustaining larval development and survival to the adult stage was also assessed. Fopius arisanus parasitized all three tephritid species, regardless of the method of exposure, but showed stronger preference for B. tryoni and B. jarvisi over B. cucumis. Superparasitism was extremely rare. Successful development of F. arisanus varied across host species. Bactrocera tryoni yielded significantly more parasitoids than B. jarvisi, but no wasps emerged from B. cucumis puparia. Tests were set up in replicated trials, but results were not homogeneous across trials. We discuss the host relationships of F. arisanus with reference to this variation and in relation to host suitability for larval development.     

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.     

Genetic manipulation of the circadian clock's timing of sexual behaviour in the Queensland fruit flies, Dacus tryoni and Dacus neohumeralis

ABSTRACT. Mating in Dacus tryoni is restricted to dusk, whereas that of a sibling species, Dacus neohumeralis , occurs in the middle of the day. The timing of sexual behaviour in both species is determined by an interaction between a circadian clock and light intensity. In D. tryoni peak mating responsiveness is at the time of dusk, and the optimal light intensity for mating is approximately 91x. In D.neohumeralis peak responsiveness is in the middle of the day, and the optimal light intensity for mating is greater than 10 000 lx. The two species were crossed and the time of mating and response to light intensity of F₁, F₂ and backcross progeny determined. The circadian clock set a mating phase ('gate') as narrow in F₁ flies as in their parents, suggesting the circadian timing mechanism to be common between the two species. The results indicate that the genetic mechanism controlling timing is independent of that controlling response to light intensity, and that both genetic mechanisms are complex.     

Winter survival and oviposition before and after overwintering of a parasitoid wasp, Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae)

Winter survival and oviposition before and after overwintering in Ooencyrtus nezarae, an egg parasitoid of phytophagous heteropterans, were examined in Osaka, Japan. Eggs of Riptortus clavatus parasitized by O. nezarae were kept under natural photoperiod and temperature. When honey was supplied, some female adults emerging from early September to late November overwintered. The percentage of overwintering individuals increased as the date of adult emergence advanced. Most female adults supplied with honey and hosts oviposited soon after emergence, then stopped laying eggs. Female adults emerging in mid‐October and early November laid eggs and then overwintered. The induction of diapause in the field seems to vary greatly depending on host availability. Without honey, the survival time of female adults was very short, whether host eggs were supplied or not. After overwintering, most females began to lay eggs in early May if host eggs were supplied, and they produced both male and female progeny. In the study area, a legume field in Osaka, parasitization by O. nezarae was observed from early July to November.     

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.     

The recovery programme for the Lord Howe Island Phasmid (Dryococelus australis) following its rediscovery

Summary   Until its rediscovery on Balls Pyramid in February 2001, the Lord Howe Island Phasmid or Stick Insect ( Dryococelus australis ) was thought to be extinct. It disappeared from Lord Howe Island soon after the accidental introduction of the Ship Rat ( Rattus rattus ) in 1918. In this paper, we report on the recovery actions undertaken for this critically endangered species since its rediscovery. Monitoring of the small surviving population on Balls Pyramid has shown it to fluctuate between about 9 and 35 adult individuals. As a safeguard against extinction, two adult pairs were removed from Balls Pyramid in February 2003 to establish captive populations in Melbourne and Sydney. Although all four founders bred readily in captivity, one pair died only a month after capture. The second female would have also died soon after capture had it not been for veterinary intervention using novel untested techniques. The single surviving pair bred successfully but the hatch rate of eggs was poor. For the next generation, both fecundity and hatch rates were low. The lack of knowledge regarding the specific husbandry requirements of this particular species undoubtedly contributed to these problems. Careful management, together with a cautious scientific approach, eventually led to all problems being resolved. Presently, there are more than 700 individuals and 14 000 eggs in captivity. Approximately 80% of incubated eggs are expected to hatch. To establish additional captive colonies, adults and eggs have been sent to other institutions, both within Australia and overseas. Now that the species is reasonably secure in captivity, the opportunity exists to reintroduce this iconic insect back onto Lord Howe Island, but this can occur only after the introduced rodents have been removed. A programme to eradicate both the Ship Rat and the House Mouse ( Mus musculus ) from Lord Howe Island is currently being developed.     

Biological parameters of Thripobius semiluteus Bouček (Hym., Eulophidae), a larval endoparasitoid of Heliothrips haemorrhoidalis (Bouché) (Thysan., Thripidae)

Abstract:  The endoparasitoid Thripobius semiluteus Bouček was recently introduced to Italy from Israel for the biological control of Heliothrips haemorrhoidalis (Bouché). In this study, some aspects of the biology of T. semiluteus were determined in the laboratory. Developmental time (egg to adult), potential fecundity, realized fecundity, progeny, daily rate of deposition of eggs and several demographic growth parameters were evaluated. Studies of the longevity of fed and starved adults at seven temperatures (3, 10, 15, 20, 23, 25 and 30 ± 1°C) showed that fed T. semiluteus lived longest at 15°C (36.45 days) and shortest at 3°C (1.45 days). Longevity was reduced significantly at higher and lower temperatures than 15°C and when honey was not provided. Development time was measured at six temperatures. Pupae did not complete development at 10°C. The lower theoretical temperature threshold was 9.0°C; the optimum developmental temperature and the upper lethal threshold were 28.0°C and 34.1°C respectively. The most rapid development was found at 30°C, but waSPS suffered significantly higher mortality at this temperature than at other temperatures. Ovigeny index showed that T. semiluteus is synovigenic, with a mean realized fecundity of 78.8 eggs per female. The mean progeny was 68.2 adults per female on fed adults.