Susceptibility of Queensland fruit fly,Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), to entomopathogenic nematodes

Queensland fruit fly, Bactrocera tryoni (Froggatt), is the economically most significant Australian tephritid pest species with a large invasion potential, yet relatively little work on its biological control has been undertaken. Entomopathogenic nematodes (EPNs) are of potential interest for control of this fruit fly species as it pupates in the soil. Specifically, the pre-pupal stage of B. tryoni may present a unique window for EPN application, as fully developed larvae drop from infested fruit to the soil for pupation. For the first time, we tested the capacity of three EPN species with different foraging strategies, Steinernema feltiae, Steinernema carpocapsae and Heterorhabditis bacteriophora, to cause larval and pupal mortality in B. tryoni across a range of EPN concentrations (50, 100, 200, 500 and 1000 infective juveniles IJs cm^-2), substrate moisture (10, 15, 20 and 25% w/v) and temperatures (15, 20, 25 and 30 °C). We found that all EPN species tested caused environment and density dependent mortality in the third larval instar while pupae were not affected. Steinernema feltiae caused high mortality across different IJ concentrations and over a wider moisture and temperature range than the other two EPN species. High mortality caused by S. carpocapsae and H. bacteriophora was more limited to high IJ concentrations and a narrower moisture and temperature range. Our findings highlight the potential of EPNs for the control of B. tryoni and warrant further laboratory and field experiments to evaluate their efficacy under the wide environmental conditions that B. tryoni can occur in.     

Sperm storage and utilization in female Queensland fruit flies (Bactrocera tryoni)

Abstract.  Female animals that use sperm from a single mating to fertilize eggs over an extended period require efficient mechanisms for sperm storage and use. There have been few studies of these mechanisms in tephritid flies. Mating, copula duration, sperm storage and sperm usage patterns are assessed in an Australian tephritid, the Queensland fruit fly ( Bactrocera tryoni ; a.k.a. 'Q-fly'). In particular, the present study investigates whether each of these aspects of mating varies in relation to female size or male size, whether sperm storage patterns change over time after mating (1, 5, 10 and 15 days), and the relative roles of the ventral receptacle and the two spermathecae as sperm storage organs. Large females are more likely to mate than are small females, and are also more fecund in the first 5 days after mating. Females are more likely to store some sperm and, among those that store some sperm, store more sperm if their mate is large. Most sperm are stored in the spermathecae (median = 97%), often with high levels of asymmetry between the two spermathecae. Asymmetry of sperm storage is related to number of sperm stored, but not to male or female size. Total number of stored sperm declines over the 15 days after mating, but this decrease in sperm numbers only reflects changes in the spermathecae; numbers of sperm in the ventral receptacle remain unchanged over this period. As a consequence, the proportion of total sperm stored in the spermathecae declines relative to the ventral receptacle. These results are consistent with a system in which small numbers of sperm are maintained in the ventral receptacle for fertilizations, and are replenished by sperm from the spermathecae as required. Sperm distribution and usage patterns in Q-flies are comparable with recent findings in medflies, Ceratitis capitata , but differ markedly from patterns found in several Anastrepha species.     

Mass-rearing and sterilisation alter mating behaviour of male Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The effects of domestication and irradiation on the mating behaviour of males of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were investigated by caging wild, laboratory-domesticated and sterile (laboratory-domesticated, gamma-irradiated) males with wild females. Mating behaviour of mass-reared males was different from that of wild males, although behaviour of wild and sterile males was similar. Mass-reared males engaged in mounting of other males much more frequently than wild and sterile males, and began calling significantly earlier before darkness. Unnatural selection pressures imposed in mass-rearing conditions may explain these changes in mass-reared male behaviour. Male calling did not appear to be associated with female choice of mating partners, although this does not exclude the possibility that calling is a cue used by females to discriminate among mating partners. Despite differences in behaviour, frequency of successful copulations and mating success were similar among wild, mass-reared and sterile males.     

柑橘大实蝇对不同柑橘品种的产卵偏好和幼虫取食选择

【目的】掌握柑橘大实蝇Bactrocera minax对不同品种柑橘的产卵偏好,以及与幼虫取食喜嗜性的关系。【方法】我们用五点取样法分别调查了自然条件下不同品种柑橘果园的虫果率,统计了各品种柑橘虫果基半部和端半部产卵痕量和卵量,并在实验室内采用培养皿桔瓣选择法研究了其幼虫对不同柑橘品种果实的取食选择。【结果】在未采取防控措施的情况下,果树面朝杂树林的方位虫果率显著高于其他方位;同一片受害柑橘园内,不同柑橘品种的虫果率为脐橙Citrus sinensis cv. Navel、酸橙C. aurantium＞冰糖橙C. sinensis cv. Bingtang、天草(Amakusa)、蜜橘C. reticulata cv. Satsuma＞沙田柚C. maxima cv. Shatian＞椪柑C. reticulata cv. Ponkan;柑橘大实蝇产卵痕数和卵量均为橙类（脐橙、酸橙和冰糖橙）大于橘类（天草、蜜橘和椪柑）和沙田柚（P＜0.05）;除蜜橘外,所调查的其他柑橘品种端半部的产卵痕数显著多于基半部（P＜0.05）;幼虫对不同品种柑橘橘瓣肉的取食选择顺序依次为：脐橙、酸橙＞冰糖橙＞天草、南橘C. reticulata cv. Nanju＞蜜橘＞椪柑＞沙田柚。【结论】结果提示,柑橘大实蝇成虫偏向于选择幼虫所喜嗜的橙类品种产卵,且产卵量更多。     

Preference and performance of Diachasmimorpha kraussii (Fullaway) (Hymenoptera: Braconidae) on five commercial fruit species

Diachasmimorpha kraussii is a polyphagous endoparasitoid of dacine fruit flies. The fruit fly hosts of D. krausii, in turn, attack a wide range of fruits and vegetables. The role that fruits play in host selection behaviour of D. kraussii has not been previously investigated. This study examines fruit preference of D. kraussii through a laboratory choice‐test trial and field fruit sampling. In the laboratory trial, oviposition preference and offspring performance measures (sex ratio, developmental time, body length, hind tibial length) of D. kraussii were investigated with respect to five fruit species [Psidium guajava L. (guava), Prunis persica L. (peach), Malus domestica Borkh. (apple), Pyrus communis L. (pear) and Citrus sinensis L. (orange)], and two fruit fly species (Bactrocera jarvisi and B. tryoni). Diachasmimorpha kraussii responded to infested fruit of all fruit types in both choice and no‐choice tests, but showed stronger preference for guava and peach in the choice tests irrespective of the species of fly larvae within the fruit. The wasp did not respond to uninfested fruit. The offspring performance measures differed in a non‐consistent fashion between the fruit types, but generally wasp offspring performed better in guava, peach and orange. The offspring sex ratio, except for one fruit/fly combination (B. jarvisi in apple), was always female biased. The combined results suggest that of the five fruits tested, guava and peach are the best fruit substrates for D. krausii. Field sampling indicated a non‐random use of available, fruit fly infested fruit by D. kraussii. Fruit fly maggots within two fruit species, Plachonia careya and Terminalia cattappa, had disproportionately higher levels of D. krausii parasitism than would be expected based on the proportion of different infested fruit species sampled, or levels of fruit fly infestation within those fruit.     

Oviposition responses of Queensland fruit fly (Bactrocera tryoni) to mineral oil deposits on tomato fruit

Behavioural responses of wild and laboratory‐culture females of Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), to mineral oil deposits on tomato fruit dipped in aqueous oil emulsions were assessed in a no‐choice test and three choice tests. The oils were two commercial products used to manage plant pests and diseases, Ampol D‐C‐Tron NR and SK EnSpray 99, one distillation fraction of the base oil of the former, and four distillation fractions of the base oil of the latter. The initial and final boiling points of the fractions were equivalent to those of n‐alkanes with chain lengths of C20–22 (Ampol), and C21–C23, C22–23.5, C22–24, and C22–24.5 (SK). For both fly types in the no‐choice test, numbers of punctures and eggs per fruit declined strongly as concentrations of the nC20–22 Ampol fraction in emulsions rose from 0.25 to 2% (vol/vol). Fly type affected the extent of responses but there was no significant interaction for fly type*oil concentration. Responses of laboratory‐culture females in the choice tests also declined as concentrations of SK and the four fractions of its base oil in emulsions rose from 0 to 0.25%. The SK nC22–24 and nC22–24.5 fractions had least impact. Responses of laboratory‐culture flies to 0.5% emulsions of the nC20–22 Ampol fraction and the nC21–23 SK fraction in choice tests were not significantly different. Likewise, responses of laboratory‐culture flies to 0.5% emulsions of the two commercial products were not significantly different. Emulsifier type did not affect numbers of punctures or eggs per fruit in choice responses of laboratory‐culture flies to 0.5% emulsions of the Ampol nC20–22 fraction or 0.5% emulsions of the SK nC21–23 fraction. If the equivalence of no‐choice and choice responses in the laboratory were to hold in the field, then unsprayed ‘sacrificial’ plants would not be necessary and oil emulsions could be used as cover sprays.     

Indirect effects of phytochemicals on offspring performance of Queensland fruit fly,Bactrocera tryoni (Diptera: Tephritidae)

Phytochemical lures such as methyl eugenol (ME) and cue‐lure are used in the management of Bactrocera fruit flies for monitoring and control. These lures are not just attractants, but also trigger physiological changes in males that lead to enhanced mating success. Additionally, in the cue‐lure‐responsive Bactrocera tryoni, females mated with lure‐fed males exhibit changes in fecundity, remating receptivity and longevity. While the lures show current generation effects, no research has been carried out on possible multigenerational effects, although such effects have been hypothesized within a ‘sexy‐son’ sexual selection model. In this study, we test for indirect, cross‐generational effects of lure exposure in F1offspring of B. tryoni females mated with cue‐lure‐fed, zingerone‐fed and lure‐unfed (=control) males. The F1 attributes we recorded were immature development time, immature survival, adult survival and adult male lure foraging. No significant differences were found between treatments for any of the three life‐history measurements, except that the offspring sired by zingerone‐fed males had a longer egg development time than cue‐lure and control offspring. However, indirect exposure to lures significantly enhanced the lure‐foraging ability of F1 adult males. More offspring of cue‐lure‐fed males arrived at a lure source in both large flight cages and small laboratory cages over a 2‐h period than did control males. The offspring of zingerone‐fed males were generally intermediate between cue‐lure and control offspring. This study provides the first evidence of a next generation effect of fruit fly male lures. While the results of this study support a ‘sexy‐son’ sexual selection mechanism for the evolution of lure response in Bactrocera fruit flies, our discussion urges caution in interpreting our results in this way.     

Effect of tomato fruit cultivar and ripening stage on Bactrocera tryoni (Froggatt) egg and larval survival

In studies of frugivorous tephritids, determining when offspring (i.e. egg and three larval instars) mortality occurs within the fruit can greatly improve the mechanistic understanding of the fly/host interaction. Previous research has demonstrated that the Queensland fruit fly, Bactrocera tryoni, has differential offspring performance in two tomato cultivars Cherry and Roma, but when juvenile mortality was occurring was not determined. We examined B. tryoni egg and larval survival in three different ripening stages (immature-green (IG), colour-break (CB) and fully-ripe (FR)) of Cherry and Roma tomato cultivars through destructive fruit sampling at 72 and 120 hr for eggs, and 48 (1st instar), 96 (2nd instar) and 120 hr (3rd instar) after fruit inoculation with neonates for larvae. Cultivar and ripening stage had no significant effect on egg survival, nor larval survival at 48 hr: the overall percentage of egg survival was at least 80% across all treatments, while 1st-instar larval was less than 52% across all treatments. In immature-green tomatoes of both varieties, nearly all mortality occurred during the first and second instars, but at 96 and 120 hr, there were significant interaction effects between cultivar and ripening stage on larval survival. In both colour-break Cherry and Roma tomatoes, there was significant larval mortality between 96 and 120 hr. However, in fully-ripe Cherry, no further significant larval mortality happened after 48 hr, while in fully-ripe Roma significant larval mortality occurred between the first and second larval instars but not thereafter. The difference in timing of larval mortality with ripening stage provides indirect evidence of active fruit defence which is strongest in immature-green fruit, less in colour-break fruit and absent in fully-ripe fruit.     

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.     

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n  = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.     

桔小实蝇和番石榴实蝇对6种寄主果实的产卵选择适应性

植食性昆虫对寄主植物的选择适应性是研究昆虫和植物协同进化关系的核心内容之一。评估寄主植物对植食性昆虫种群的适合度,需要综合分析昆虫对寄主的产卵选择性和寄主对昆虫的取食适合性。以桔小实蝇和番石榴实蝇为研究对象,分别测定了这两种实蝇对6种寄主果实:番石榴、香蕉、杨桃、木瓜、甜橙、番茄的产卵选择性以及幼虫取食后对其生长发育的影响。寄主产卵选择性实验分别采用完整寄主果实直接供试产卵和块状寄主果实气味引诱产卵两种处理方式;在生长发育适应性实验中,以幼虫和蛹的存活和生长发育等相关参数作为评价指标。实验结果表明,寄主的供试方式不同,两种实蝇的选择性均有明显差异;对寄主气味选择性强的寄主更适合于两种实蝇后代的生长发育。两种实蝇对6种寄主果实的产卵选择性和后代发育适合性两者相关性不显著,与许多文献报道单一地采用发育适合性(如发育历期、存活率或蛹重等)作为评价寄主选择性的结果不一致。两种实蝇之间对6种寄主果实的产卵选择和幼虫取食适合性既具相似性也具差异性,表明这两种实蝇在寄主生态位上既有重叠性又有分化性。     

Feeding behaviour of Bactrocera cacuminata (Hering) (Diptera: Tephritidae) on methyl eugenol: a laboratory assay

Abstract Many dacine (Diptera: Tephritidae) species are attracted to one of two chemical substances, 4-allyl-1,2-dimethoxybenzene (methyl eugenol (ME)) or 4-( p -hydroxyphenyl)-2-butanone (cue lure). Despite the fact that these chemicals or analogs occur naturally, their significance in the biology/ecology of the Dacinae has seldom been examined. In this study, we examine the patterns in feeding behaviour of Bactrocera cacuminata (Hering), a monophagous non-pest fruit fly, on ME. Based on a laboratory assay, we found that males of this species fed on ME multiple times within a single day and on multiple days. The pattern of repeat feeding was not related to time since previous feeding or duration of prior feeding. Our results contrast those obtained on a similar study on a related polyphagous species ( Bactrocera dorsalis (Hendel)). We discuss the implications of our findings with a view to explaining the functional significance of dacine response to these plant-derived chemicals.     

Behavioural evidence for hygro- and mechanoreception by ovipositor sensilla of Dacus tryoni (Diptera: Tephritidae)

ABSTRACT. Gravid female Queensland fruit flies Dacus tryoni (Froggatt) readily oviposit when the ovipositor sensilla are stimulated either by a relatively thick (≥2 mm) surface layer, or by moisture under a relatively thin (<0.5 mm) or thick surface layer. A single layer of Parafilm over dry air elicits almost no oviposition; significantly more oviposition occurs when the underside of the Parafilm is coated with a 2 mm thick layer of petroleum jelly; even more oviposition occurs when the Parafilm is floated on liquid paraffin; greatest oviposition is elicited when moist media (agar gel, water or moist air) are presented under the Parafilm. Hollowed-out domes of apple skin elicit significant oviposition only when sealed basally, to give high internal concentrations of water vapour. The previously identified groove sensilla on the tephritid ovipositor are inferred as being a source of mechanical and chemical information used by the fly in such ovipositional decisions. The ovipositor also has two other types of tactile sensilla. The three types of ovipositor sensilla of tephritids are considered to play roles analogous to those of certain labial receptors of flies with piercing mouthparts.     

Ant cues affect the oviposition behaviour of fruit flies (Diptera: Tephritidae) in Africa

Abstract. Although most studies on fruit fly oviposition behaviour focus on horizontal interactions with competitors and cues from host plants, vertical interactions with predators are poorly documented. The present study provides direct evidence indicating that the oviposition behaviour of the two main mango fruit fly species, Ceratitis cosyra (Walker) and Bactrocera invadens Drew‐Tsurata & White, is affected by secretions of the dominant arboreal ant Oecophylla longinoda (Latreille). When offered ant‐exposed and unexposed mangoes in the absence of the ants, both fly species are reluctant to land on ant‐exposed fruits and, when having landed, often take off quickly and fail to oviposit. The number of puparia collected from unexposed mangoes is approximately eight‐fold higher than from ant‐exposed ones. The results obtained from laboratory experiments and field observations confirm that adult fruit flies are more affected through repellence by ant cues than by direct predation. The use of cues by fruit flies in predator avoidance has implications for evolutionary ecology, behavioural ecology and chemical ecology.