Ant-coccid mutualism in citrus canopies and its effect on natural enemies of red scale, Aonidiella aurantii (Maskell) (Hemiptera: Diaspididae).

Mutualistic relationships between honeydew-producing insects and ants have been widely recognized for several decades. Iridomyrmex rufoniger (Lowne) is the commonest ant species associated with black scale, Saissetia oleae (Olivier), in the citrus orchards of the mid latitudes of coastal New South Wales. Citrus trees with high densities of both red and black scale and high ant activity were identified and the results of excluding ants from half of those trees (using a polybutene band on each trunk) were compared with the results of not excluding ants from the other half. Trees with a low incidence of black scale and ants were also studied. Exclusion of ants from trees was soon followed by collapse of black scale populations because most individuals were asphyxiated by their own honeydew. Also, parasitism of the red scale by Encarsia perniciosi (Tower) and Encarsia citrina Craw was significantly higher than in the control trees over the following year, as was the predation rate on red scale due to three coccinellid predators, Halmus chalybeus (Boisduval), Rhyzobius hirtellus Crotch and Rhyzobius lophanthae (Blaisdell). In contrast, another coccinellid, Orcus australasiae (Boisduval), and a noctuid moth larva, Mataeomera dubia Butler, were seen in low numbers on banded (ant exclusion) trees, probably because of the low availability of their black scale prey, but were significantly higher on control trees apparently because of their invulnerability to ants.

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.

Multiple mating by females of two Bactrocera species (Diptera: Tephritidae: Dacinae).

Multiple mating was investigated in two tephritid species when females were under minimal male pressure because they were each confined with a single male in cages 20 x 20 x 20 cm and observed daily until they died. Laboratory-reared females of Bactrocera cucumis (French) lived up to 274 days and refractory periods averaged 59-63 days. However, the distribution of matings among B. cucumis females was not significantly different to that expected by chance. Wild females of Bactrocera cacuminata (Hering) reared from field-collected fruits of Solanum mauritianum Scopoli lived for up to 134 days and mated up to three times with refractory periods between matings averaging 27-39 days. The distribution of the number of matings among females of B. cacuminata was non-random because of the high proportion of non-maters (50%); but, when only females mating more than once were considered, there was no significant departure from random expectation.

The genetic structure of populations of an invading pest fruit fly, Bactrocera tryoni, at the species climatic range limit.

Previous population genetic studies of the Queensland fruit fly, Bactrocera tryoni Froggatt (Diptera: Tephritidae), in its central range have shown barely detectable genetic differentiation across distances of almost 3000 km (F(st)=0.003). In this study, we investigated the genetic structuring of southern border populations of B. tryoni, in the region extending from the central population to the recently colonized southern range limit. The expectation was that marginal populations would be small, fragmented population sinks, with local adaptation limited by gene flow or drift. Unexpectedly, we found that the population at the southern extreme of the range was a source population, rather than a sink, for the surrounding region. This was shown by assignment testing of recent outbreaks in an adjoining quarantine area and by indirect migration estimates. Furthermore, populations in the region had formed a latitudinal cline in microsatellite allele frequencies, spanning the region between the central population and the southern range limit. The cline has formed within 250 generations of the initial invasion and appears stable between years. We show that there is restricted gene flow in the region and that effective population sizes are of the order of 10(2)-10(3). Although the cline may result from natural selection, neutral evolutionary processes may also explain our findings.

Laboratory adaptation of Bactrocera tryoni (Diptera: Tephritidae) decreases mating age and increases protein consumption and number of eggs produced per milligram of protein.

A significant reduction in age of mating occurred during the first four generations (G1-G4) of laboratory adaptation of wild Bactrocera tryoni (Froggatt) and this was associated with the earlier attainment of peak egg load although no significant differences were detected in the peak egg load itself. A long term laboratory (LTL) strain had a significantly earlier mating age and higher peak egg load than flies of wild origin or those from the first four laboratory generations. The amount of protein consumed by females in the first week of adult life was significantly higher in the LTL strain than in flies of wild origin or G1-G4 but there were no significant changes (or only slight changes) with laboratory adaptation in the amounts of protein consumed up to the ages of mating and peak egg load. Laboratory adaptation resulted in no significant changes in egg size, egg dry weight, puparial fresh weight and the dry weight of newly emerged females. The large increase in fecundity with laboratory adaptation is associated with a 4- to 5-fold increase in the rate of conversion of dietary protein to eggs (i.e. eggs produced per mg of protein consumed).

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.

Towards a male-only release system for SIT with the Queensland fruit fly, Bactrocera tryoni, using a genetic sexing strain with a temperature-sensitive lethal mutation.

Flies that are homozygous for the recessive autosomal mutation bent wings have a limited ability to fly and are less tolerant of high temperatures than normal flies in both the egg and puparial stages. The differences between the mutant and normal flies were found sufficient to be the basis of a genetic sexing strain. Genetic sexing strains were created using translocations of the autosome bearing the wild-type allele of bent wings (chromosome 2) to the Y chromosome, and crossing male flies carrying the translocation to mutant bent wings females. In the resulting strain, the females were homozygous for the bent wings mutation and the males were phenotypically normal for wing characters. Several translocations were recovered after irradiation, but only one translocation involving chromosome 2 was both stable and expressed in a stock that was vigorous enough for long-term viability. Unfortunately, all stocks containing the translocation showed high levels of temperature-dependent lethality, including, inexplicably, both males and females. Translocation stocks showing this effect included bent wings, another second chromosome mutation, white marks, and an otherwise normal stock. This phenomenon is probably rare, as it has not been reported before. It is likely that bent wings could be suitably used with another translocation.

Integrated pest management of two-spotted mite Tetranychus urticae on greenhouse roses using petroleum spray oil and the predatory mite Phytoseiulus persimilis.

From 1995 to 1999, four experiments were conducted on greenhouse roses to assess the effectiveness of the nC24 petroleum spray oil (PSO), D-C-Tron Plus, against two-spotted mite, Tetranychus urticae Koch (Acarina: Tetranychidae), and to determine how the oil could be most efficiently and effectively used in combination with the predatory mite Phytoseiulus persimilis Athias-Henriot (Acarina: Phytoseiidae) in an integrated pest management program. The results showed that 0.5% PSO applied fortnightly to roses gave excellent protection from T urticae infestation when the mite population was not already established. However, PSO applied after roses were infested with T. urticae above the economic threshold only stabilised populations without reducing them below that threshold. Populations of P. persimilis in the upper and lower canopies were unchanged after two sprays of PSO at 7-day intervals, and application of PSO to the upper canopy was as effective in controlling T. urticae in the presence of P persimilis as spraying the entire plant. Combining PSO with P. persimilis gave better control of T. urticae than using P. persimilis alone. The most cost-effective use of PSO in the presence of P. persimilis is, therefore, to apply spray only to the upper canopy. This will not affect control of powdery mildew with PSO. Comparison of a control program for T urticae based on the monitored use of synthetic miticides with that based on calendar application of PSO revealed that both gave equally effective control. The benefits of combining PSO and P. persimilis in an integrated pest management program for T. urticae on roses over a program based on synthetic fungicides are discussed.

Microsatellite analysis of the Queensland fruit fly Bactrocera tryoni (Diptera: Tephritidae) indicates spatial structuring: implications for population control.

The population structure of a tephritid pest species, the Queensland fruit fly Bactrocera tryoni (Froggatt), has been analysed over a five year period (1994-1998), using six microsatellites. Adult fly samples were collected to cover most regions of eastern and central Australia where the flies are regularly found. Tests for heterogeneity indicated that flies within geographically defined regions were homogeneous. The samples were allocated into five regions, including one very large region, Queensland, which encompasses that portion of the fly's range where breeding can occur year-round. With one exception, the collections from different regions were homogeneous between years, showing a fairly static distribution of the species. However, differences between regions were highly significant. The one case of a change in frequency between years indicated a gradual replacement of flies in a marginal region by flies from the main part of the range. The finding of stability in the distribution of a highly mobile insect is of interest, potentially also for other species which have expanded beyond their native range. It is argued that a contributing reason for this stability may be adaptation to different climatic regimes, and that strategies for control based on this hypothesis afford a reasonable chance of success.

Seasonal trends in acclimatization to cold in the Queensland fruit fly (Dacus tryoni, Diptera) and their prediction by means of a physiological model fed with climatological data.

This paper demonstrates the predictability and significance of changes in the temperature threshold for cold-torpor and in the threshold for survival at sub-zero temperatures in two contrasting types of climate. Both thresholds alter on a seasonal basis in both types of climate and it is observed that temperatures prevailing in autumn cause acclimatization for winter long before winter arrives.Torpor thresholds can be accurately predicted in laboratory-simulated climates by use of a model. When the model is used for field predictions, using shade temperature data from the field, it is only successful in predicting accurate threshold values for those parts of the year during which there is a downward trend in threshold.The relative importance of torpor-avoidance and frost-resistance is discussed with reference to records of the frequency and extent of various degrees of cold in the field. It is concluded that the marked lowering of torpor threshold prior to winter is of great significance in enabling overwintering flies to stay in their sheltered hibernacula. The relatively slight lowering of temperature threshold for frost-survival is only of significance in certain climates. It greatly improves chances of forst-survival in climates where yearly minima fall predominantly in the range-2.65°C to-4.05°C. Acclimation is not necessary for frost survival in climates where the yearly minima stay above-2.65°C whereas acclimation is ineffective in preventing frostinduced extinction when yearly minima fall below-4.5°C.

The developmental dynamics of Tipula paludosa and the relation of climate to its growth pattern, flight season and geographical distribution.

The model of Meats (1974a) is tested against Laughlin's (1967a) data on growth in the field. A good measure of agreement is found. The model is then used to investigate the consequences of the univoltine T. paludosa starting its life cycle (flight season) at various dates in the range quoted by Coulson (1962)-i.e. from July to October. Similarly the influence of increased or decreased monthly temperatures is also investigated. The model indicates that growth will be poorer the later the life cycle starts or the lower the mean monthly temperature.Populations at Moor House (high Pennines) are probably obliged to start their cycle earlier by shortening the prepupal/pupal period because the relatively colder average temperatures would otherwise prevent the attainment of a suitably sized adult. Lowland populations from Canada, England, Germany and Poland differ in developmental schedule depending on prevailing temperatures but usually achieve sufficient growth despite a relatively prolonged prepupal/pupal period and late flight season. Dry conditions (which, like low temperatures, can slow growth) are not associated with advanced flight seasons. In areas where low temperatures are critical, successful growth may be restricted to wet soils. The relation of climate to prepupal diapause is discussed and developmental dynamics is related to potential geographical range.

A population model for two species of Tipula (Diptera, Nematocera) derived from data on their physiological relations with their environment.

Published results show that variations in mortality due to factors such as predation, parasitism and disease are overshadowed by variations due to effects of physical factors. Soil temperature and the tension of the soil moisture have a profound effect not only on the survival but also on the growth rate and fertility of the two species considered here. The model is based largely upon known and experimentally quantified relationships of causes and effects and is therefore a practical model which integrates our knowledge so that we can compare its results with reality to see if or where our knowledge is lacking.

Simulation of population trends of Tipula paludosa using a model fed with climatological data.

This paper tests the model of Meats (1974) by comparing its simulations with the real data of Milne et al. (1965) and of Laughlin (1967). Simulations are also made for conditions not previously investigated.

The relative importance to population increase of fluctuations in mortality, fecundity and the time variables of the reproductive schedule.

Previous authors have used simple models to investigate the relative importance to population increase of variations in the total and age-specific reproductive rates. But while acknowledging that the latter were the product of the age specific birth and death rates, they have used their models only to investigate changes in total or age-specific birth rates and have not been concerned with variations in death rates. This paper extends the use of Lewontin's (1965) model, to a wide range of values of r, the exponential rate of population increase. It shows how the relative importance of changes in certain life-history features can change with r and be reversed when r is near to zero. It is also shown that variations in mortality rate are not necessarily best expressed in analogous terms to variations in birth rate. If more suitable terms are used it is seen that changes in mortality rate can be of varying importance depending on the existing mortality rate. They can be overwhelmingly important when the mortality rate is high.