Behavioural responses of female Queensland fruit fly, Bactrocera tryoni, to mineral oil deposits

Behavioural responses of Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), females to fruit dipped in water and fruit dipped in 0.5% (vol/vol) aqueous emulsions of a mineral oil were determined and analysed. The mineral oil was an nC20–22 distillation fraction of the base oil used to produce an nC23 horticultural mineral oil. Females caged with oil‐treated fruit had significantly longer prelanding intervals than females caged with water‐dipped fruit. The latter was attacked immediately or shortly after being caged with flies whereas some oil‐dipped fruit was not attacked within 180 min. The percentage of landings that led to oviposition on water‐ and oil‐treated fruit were 58 and 13%, respectively, and the percentages ovipositing after probing were 74 and 25%, respectively. Likewise, average times spent probing were 7 vs. 31 s whereas average times spent ovipositing were 321 vs. 223 s. Females spent less than half as much time on oil‐treated fruit than on water‐treated fruit. Transition probabilities of rejection, when applied to the behaviour sequence indicated that oil‐treated fruits are about nine times less likely to be infested with B. tryoni.     

矿物油乳剂拒避橘小实蝇的有效组分分析

高品质的矿物油乳剂可用于有机食品生产,在害虫综合治理中有良好的应用前景。矿物油是包含多种化学成分的混合物,其组成成分和比例决定其理化性质并关系到其防治病虫害的效果。已有研究报道表明,矿物油乳剂对许多害虫有拒避作用,但其机理及相关性组成成分尚未研究清楚。本实验测定了4种矿物油乳剂即SK绿颖农用喷洒油、加德士-路易夏用油、安波尔喷洒油、法夏乐石蜡油（分别简称为SK,Caltex,Ampol和Citrole油）和Caltex油＜120℃,＞120℃,＜105℃,105~120℃及SK油＜110℃、＞110℃等6种分子蒸馏组分的正构烷烃碳数当量nCy值、碳数分布值及其对橘小实蝇Bactrocera dorsalis的产卵拒避效果。室内生物测定结果显示,橘小实蝇在SK,Caltex,Ampol和Citrole油0.5%水乳液及清水浸蘸处理后的香蕉上平均产卵孔数分别为17.850±1.282,3.400±0.630,15.850±1.439,20.650±1.212和19.050±1.155,在同样方法处理的阳桃上平均产卵孔数分别为15.500±3.969,3.000±0.707,13.750±4.131,17.250±4.385和62.000±4.708。橘小实蝇在6种分子蒸馏组分0.5%水乳液处理后的香蕉上的平均产卵孔数分别为4.400±1.166,8.200±0.583,5.400±1.720,2.400±1.077,4.800±1.594和12.200±2.63,显示不同矿物油的拒避效果差异显著（P<0.05）,拒避效果最好的矿物油的正构烷烃碳数当量nCy值在nC23左右.矿物油对橘小实蝇的产卵拒避效果与其相应的碳数分布值的Spearman等级相关性分析显示,其拒避效果与矿物油中的C23组分的含量呈显著正相关关系（P<0.05）。这可为进一步分析矿物油乳剂拒避害虫的作用成分和探索其机理提供参考。     

矿物油乳剂作用下橘小实蝇的产卵拒避及触角电位反应

为了研究矿物油乳剂对害虫行为的干扰作用,测定和比较了4种矿物油乳剂对橘小实蝇Bactrocera dorsalis (Hendel)的产卵拒避效果及在其作用下该虫的触角电位反应（EAG）,并测试了其中拒避效果最好的Caltex Lovis夏用油的不同施用方式（浸蘸法和喷雾法）及不同施用浓度（150,250,350,450倍液）对橘小实蝇产卵拒避的影响。结果显示：分别用4种矿物油乳剂200倍液以浸蘸法处理香蕉,仅Caltex Lovis夏用油处理对橘小实蝇产卵拒避效果显著,其平均产卵孔数仅为清水对照的15.14%;其他3种矿物油乳剂则没有明显效果。这表明矿物油乳剂对昆虫行为的干扰作用与其组成成分相关。4种矿物油乳剂对橘小实蝇的产卵拒避效果与各矿物油乳剂的50倍液滴于滤纸条后0.5 h的雌成虫EAG值间无显著相关性,但与滴于滤纸条后3 h（滤纸条上水分已蒸发）的EAG值间有显著的负相关性,说明矿物油乳剂的挥发性组分对橘小实蝇的产卵拒避影响不大,因此推测矿物油乳剂在植物表面形成的油膜对植物挥发性物质的封闭作用与其产卵拒避效果可能有一定的关系。浸蘸法不同浓度处理间的产卵拒避效果无显著差异,均明显高于喷雾法处理及清水对照。在喷雾法处理中,仅150倍液有显著产卵拒避效果。这说明,矿物油乳剂需要在植物表面达到一定的沉积量,才能对橘小实蝇有较好的产卵拒避作用。与喷雾法相比,浸蘸法可能较利于矿物油在植物表面的沉积,这也佐证了矿物油乳剂所形成油膜的封闭性与其产卵拒避效果有一定的关系。     

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.     

Predation potential of the earwig Euborellia annulipes on fruit fly larvae and trophic interactions with the parasitoid Diachasmimorpha longicaudata

The earwig Euborellia annulipes (Lucas) (Dermaptera: Anisolabididae), a generalist predator, has been observed in fruits infested with fruit fly larvae, which are frequently parasitized by parasitoid wasps. Neither the capacity of earwigs to predate on fruit flies nor intraguild interactions between earwigs and fruit fly parasitoids have been investigated. Here, we studied in laboratory conditions the predation on the fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) by the earwig E. annulipes, and whether parasitism of fruit fly larvae by the parasitoid wasp Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) influences predation by the earwig. We evaluated the predation capacity, functional response and prey preference of E. annulipes for parasitized and non-parasitized fruit fly larvae in choice and no-choice tests. We found that earwigs prey on second- and third-instar larvae and pupae of C. capitata and consumed larger numbers of second-instar larvae, followed by third-instar larvae and pupae. Females prey on larger numbers of fruit flies than did males, regardless of the prey developmental stage, but both sexes exhibited a type II functional response. Interestingly, males killed but did not consume fruit fly larvae more than did females. In no-choice tests, earwig females consumed equal numbers of parasitized and non-parasitized fruit fly larvae. However, in choice tests, the females avoided feeding on parasitized larvae. Subsequent tests with hexane-washed parasitized and non-parasitized larvae showed that putative chemical markings left on fruit flies by parasitoids did not drive the earwig preference towards non-parasitized larvae. These findings suggest that E. annulipes is a potential biological control agent for C. capitata, and that, because the earwig avoids consuming larvae parasitized by D. longicaudata, a combination of the two natural enemies could have an additive effect on pest mortality.     

Mechanism and effectiveness of safflower oil against female Queensland fruit fly Bactrocera tryoni

Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), infests many horticultural fruit crops in the eastern part of Australia. Farmers usually apply synthetic insecticides to control this pest. Little is known on the use of plant products especially vegetable oils for fruit fly control although they are considered to be safer than synthetic insecticides. In this study, safflower oil was investigated for its mechanism and effectiveness against female B. tryoni. In a laboratory test, safflower oil treatments (2.5 and 5.0 ml l^?1) reduced the number of fly punctures on treated artificial fruits, no matter whether pre‐punctures were present or absent. Safflower oil treatments also reduced the number of fly landings and eggs laid, but only when the treated artificial fruits were without pre‐punctures. These results confirmed that safflower oil is active against female B. tryoni mainly by preventing this fruit fly from making oviposition punctures, not by discouraging them from depositing eggs or by repelling them. The slippery nature of safflower oil is considered to be responsible for a reduction in the susceptibility of artificial fruit to fruit fly punctures. Further investigation using fruit‐bearing tomato plants (a no‐choice test) in a glasshouse situation revealed that safflower oil application at concentrations of 10 and 15 ml l^?1 reduced the number of oviposition punctures but failed to reduce the number of eggs laid. To increase efficacy of safflower oil under field conditions, multiple tools may be needed to reduce fruit fly populations and oviposition behaviour, such as the addition of trap‐crops, provision of artificial oviposition sites, or mixing the insecticides with the oil.     

Effect of food deprivation on attractiveness of food sources, containing natural and artificial sugar and protein, to three African fruit flies: Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis capitata

The effects of food deprivation, age, and mating status on the responses of three fruit fly species, Ceratitis cosyra (Walker), Ceratitits fasciventris (Bezzi), and Ceratitits capitata (Wiedemann) (Diptera: Tephritidae) to natural and artificial sugar and protein food sources were investigated. Natural food sources included guava [Psidium guajava L. (Myrtaceae)] juice (a common host fruit for all three fruit fly species) and bird faeces (farm chicken). Artificial food sources included molasses (obtained from a local sugar factory) and a locally produced protein bait (the International Centre of Insect Physiology and Ecology yeast). In all species studied, sugar deprivation of immature (1–2‐day‐old) male and female flies increased their response to food odours, although it did not change their preference for the type of odour (protein or sugar). Protein deprivation of mature (14–17‐day‐old) male and female flies also increased their response to food odours compared to protein‐fed flies. Protein‐deprived females were highly attracted to odours from protein sources in particular. Odours from natural food sources, guava juice, and chicken faeces, were more attractive to food‐deprived flies than were odours from artificial sugar and protein sources. Attraction to food odours increased significantly with increasing age for protein‐deprived females of all species. For males and females of all species, nutritional state was a more important factor than mating status in influencing responses of flies to food odours. Practical implications of these findings are discussed in terms of strategies for fruit fly control using food baits.     

Electroantennogram response and attraction of Anastrepha suspensa to volatiles of various sugar sources and aged sugar solutions

With the aim of finding new, sugar‐based volatile attractants for economically important tephritid fruit fly species, we used electroantennography (EAG) to quantitate olfactory responses of female Caribbean fruit fly, Anastrepha suspensa (Loew) (Diptera: Tephritidae), to volatiles of six sugar sources (refined white and brown cane sugar, coconut sugar, date sugar, date jaggery, and cane panela). Laboratory‐strain and wild flies, both sexually immature and mature, were tested for EAG responses to the volatiles of dry crystallized sugar sources and 10% (wt/vol) aqueous solutions that had aged in the laboratory for 0–7 days. In general, wild flies exhibited higher EAG responses than laboratory flies, and immature females responded more strongly than mature females. With the exception of date jaggery and cane panela, volatiles of dry sugar sources and 0‐ and 1‐day‐old solutions elicited lower EAG responses than any of the aged solutions. Most solution volatiles elicited the strongest EAG response after 2 days of aging. Of the treatments evaluated, volatiles of the 5‐day‐old date jaggery solution elicited the highest‐amplitude EAG responses (39%) in A. suspensa females. On the basis of the latter, we tested the attraction response of mature and immature females to date‐jaggery solutions aged over 2 and 4 days in two‐choice flight tunnel bioassays. With both mature and immature females, the 2‐day‐old solution was more attractive than the 4‐day‐old jaggery solution, but significantly more mature females (70% of captures) were attracted to 2‐day‐old jaggery solution. We discuss our results with respect to the improvement of fruit fly lures and attractants by incorporating elements from aged date‐jaggery sugar.     

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.     

Oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens

We assessed the role of visual and olfactory cues on oviposition preference in the oligophagous tomato fruit fly, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae). In a field survey, we evaluated the stage of susceptibility of field‐grown tomatoes by monitoring N. cyanescens infestations from fruit‐setting up to harvest, in relation to post‐flowering time, size, and visual properties of fruit. In two‐choice laboratory experiments, we tested the degree to which females use visual and olfactory cues to select their host plant for oviposition. In addition, we investigated the ability of flies to avoid fruit already infested by conspecific eggs or larvae, and the influence of natal host fruit on oviposition preference. Neoceratitis cyanescens females preferentially lay their eggs in small yellow‐green unripe fruit (2–3.5 cm diameter, 10–21 days post‐flowering). Damage to fruit was significantly affected by brightness and size properties. In laboratory experiments, females chose to lay their eggs in bright orange rather than yellow domes. On the sole basis of olfactory stimuli, females showed a significant preference for unripe vs. ripe host fruit, for unripe fruit vs. flowers or leaves, and for host vs. non‐host fruit (or control). However, colour interacted with odour as females dispatched their eggs equally between the yellow dome and the bright orange dome when unripe fruit of tomato was placed under the yellow dome vs. ripe fruit under the bright orange dome. When offered real ripe and unripe tomatoes, females preferred unripe tomatoes. Females significantly chose to lay eggs in non‐infested fruit when they were given the choice between these or fruit infested with larvae. In contrast, recent stings containing eggs did not deter females from laying eggs. Rather, they could have an attractive effect when deposited within <1 h. Regardless of their natal host plant, tomato or bugweed, N. cyanescens females laid significantly more eggs in a dome containing bugweed fruit. However, 15% of females originating from tomato laid eggs exclusively in the dome with tomato, against 3% of females originating from bugweed.     

Essential oils of citrus fruit stimulate oviposition in the Mediterranean fruit fly Ceratitis capitata (Diptera: Tephritidae)

Female Mediterranean fruit flies (medfly) Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) perceive both qualitative and quantitative aspects of citrus fruit chemistry. However, the behavioural and biological adjustments of this response remain largely unknown. In the present study, the ovipositional responses of gravid female medflies to essential oils (i.e. the most critical resistance factor to medfly infestation) of various citrus fruits are investigated. In dual‐choice (treatment versus distilled water control) experiments, females lay significantly more eggs into pre‐punctured hollow oviposition hemispheres (domes) provided with 1 µL of citrus peel oil from sweet orange, satsuma mandarin, bitter orange, grapefruit and lemon compared with odourless domes. No‐choice tests show a weak effect of lemon essential oils in stimulating oviposition. The female ovipositional response to sweet orange oil (the most active in eliciting oviposition) is dose‐dependent. Additionally, limonene, the most abundant chemical in all citrus oils, stimulates oviposition, whereas linalool, a representative compound of immature citrus fruit associated with high toxicity against immature stages of fruit flies, has a significant deterrent effect. In further no‐choice tests, females lay approximately 23% fewer eggs in limonene (93%) (amount found in orange oil) and 60% fewer eggs in limonene 93% plus linalool 3% (approximately 10‐fold the amount found in orange oil) mixtures, relative to sweet orange oil. The results suggest that the limonene content accounts largely (but not completely) for the ovipositional responses observed in sweet orange oil, whereas high linalool proportions are capable of significantly masking and/or disrupting its stimulatory effects in citrus oils. The importance and practical implications of these findings with respect to understanding how citrus fruit chemistry influences the ovipositional responses of medfly is discussed.     

Flower and fruit volatiles assist host‐plant location in the Tomato fruit fly Neoceratitis cyanescens

Mature females of the tomato fruit fly Neoceratitis cyanescens can detect host fruit at a short distance using only visual stimuli, but little is known about the role of airborne volatile cues in the host searching strategy. A series of experiments is conducted in a laboratory wind tunnel, in which the behavioural responses of individual flies to volatiles from Solanaceae host plants (including tomato Lycopersicum esculentum Mill., bug weed Solanum mauritianum Scop. and Turkey berry Solanum torvum Sw.) are observed, according to some environmental (air speed) and physiological (age and mating status of females, time of day) factors. Mature females respond primarily to specific olfactory cues from blends of flowers or host fruit, preferentially unripe fruit for bug weed, as opposed to ripe fruit for Turkey berry or tomato. Males are also highly attracted by the odour of unripe fruit of bug weed. Wind plays a key role, as shown by the proportion of flies that reach the upwind section of the tunnel in the presence of both fruit odour and air flow (66.7%) and in the absence of either fruit odour (13.3%) or wind (36.7%). In response to fruit volatiles carried by wind, flies embark in a ‘plume tracking’ or ‘aim and shoot' flight, consistent with odour‐conditioned anemotaxis. Females respond to host fruit odour regardless of their age, egg load or mating status, and also more consistently in the afternoon, which is their preferential time of day for egg‐laying. Searching behaviour and response to host volatiles in N. cyanescens are discussed in the light of host‐finding and an adaptive strategy.     

Food deprivation effects on carbohydrate levels and their relation to feeding responses of western cherry fruit fly Rhagoletis indifferens to spinosad bait

Abstract The nutritional state of tephritid fruit flies affects various behaviours. The present study aims to determine food deprivation effects on carbohydrate levels and their relation to feeding responses to spinosad bait (GF‐120^® Naturalyte^® Fruit Fly Bait; Dow AgroSciences, Indianapolis, Indiana), as measured indirectly by mortality, in western cherry fruit fly Rhagoletis indifferens Curran (Diptera: Tephritidae). Sugar levels in 1–2‐day‐old flies exposed to sugar for 1 h and then deprived of sugar for 10–24 h decrease but, in flies with access to continuous sugar and no sugar, they increase and do not change, respectively. Sugar levels in 14–15‐day‐old flies that have had free access to yeast extract and sugar and are then deprived of it for 0 and 10 h do not differ but they are lower at 24 and 30 h. Mortalities of 1–2‐day‐old flies exposed to sugar for 1 h and then deprived of it for 0–24 h progressively increase but they do not increase in flies given no sugar because the mortalities in this treatment are equally high at all times. By contrast, mortalities of 14–15‐day‐old flies deprived of food increase from 0 to 10–30 h but there are no differences from 10 to 30 h. Mortalities of 14–15‐day‐old flies deprived of food for 3 and 6 h also do not differ. Food deprivation effects on glycogen are similar, although glycogen occurs at lower levels than sugar. The results obtained suggest that R. indifferens flies need to feed multiple times on carbohydrate foods during the day to maintain their carbohydrate levels, that their responses to spinosad bait will increase if they do not, and that there may be age‐related effects on carbohydrate levels and responses to spinosad bait.     

Combination of edible vegetable oil and artificial fruit to reduce Bactrocera dorsalis oviposition in chilli fruits

Bactrocera dorsalis (Hendel) is a major pest of a wide variety of fruits and vegetables. This study aimed to evaluate the potency of combining edible vegetable oil application and artificial fruit provision to reduce the oviposition of B. dorsalis in chilli fruits. Experiments were conducted in laboratory using a 20‐L plastic container provided with chilli fruits and in semi field using a fruit‐bearing chilli plant caged with insect screen. The laboratory test revealed that the combination of coconut oil application and artificial fruit provision significantly reduced fruit fly visits, eggs laid and the number of infested chilli fruits no matter when the chilli fruits were without prepuncture or with prepuncture. Further study using a caged chilli plant also found that this combination had significant effects on fruit fly visits and infestation in chilli fruits. These results provide an indication that coconut oil and artificial fruit can work together to reduce fruit fly attacks on chilli fruits. Combining the use of coconut oil and artificial fruit is likely to create an integrated behavioural manipulation (push and pull) of female B. dorsalis that lead to a much lower fruit fly infestation in chilli fruits.     

Determination of volatile substances in olives and their effect on reproduction of the olive fruit fly

Females of the olive fruit fly Bactrocera oleae lay their eggs in olives mainly using fruit volatile stimuli. Using GC-MS analysis, we determined the chemical composition of the volatile blend emitted from field-collected olive fruit of cv. Megaritiki, at different stages of maturity. GC-MS analysis demonstrated qualitative and quantitative differences in the headspace blend emitted by the olive fruit. Certain chemicals such as toluene, n-octane, α-pinene, limonene, ethyl hexanol, nonanal n-dodecane, decanal and n-tetradecane were detected in greater amounts, irrespective of the growth stage of the fruit. The flies’ exposure to a number of these chemicals, such as n-octane and α-pinene, as well as to a mixture consisting of n-octane, α-pinene, limonene, ethyl hexanol, nonanal, n-dodecane, decanal and n–tetradecane favoured successful mating and egg production. The results may contribute to the improvement of the mass rearing of the fly, through these findings, which illustrate the positive effect that certain fruit volatile chemicals have on the fly's reproduction and offer a better understanding of the relation between the fly and the host fruit.     

Visual response of the tomato fruit fly, Neoceratitis cyanescens, to colored fruit models

To determine how mature females of the tomato fruit fly, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae), detect host fruit after arriving in their host plant habitat, behavioral responses to colored models were observed in a laboratory flight chamber. Host‐seeking females oriented themselves preferentially towards bright orange spheres (3.7 cm in diameter), irrespective of their natal host fruit: tomato, bug weed, or black nightshade. Females oriented themselves preferentially towards the orange sphere when placed against a fluorescent yellow background as opposed to a black background, but the distribution of responses to the set of colored spheres did not vary significantly with background color. In a choice situation between bright orange spheres of various sizes (1.9, 3.7, and 7.5 cm in diameter), females landed preferentially on the bigger sphere. However, they preferred a yellow color when the latter was associated with two‐dimensional models, probably mimicking leaves. The attractiveness of orange spheres depended more on the proportion of reflected light in the spectral region around 610 nm than brightness of color in itself. Low light intensity significantly influenced the activity of the flies but not their visual preference. The strong response of females to bright orange spheres confirmed the importance of visual characteristics in short‐range mechanisms of host‐plant location in specialist insects. Responses to fruit visual stimuli are discussed relative to other Tephritidae, host‐finding strategy, and pest management.     

Evidence for sexual isolation as a prezygotic barrier to gene flow between morphologically divergent species of Rhagoletis fruit flies

1. Certain groups of fruit flies in the genus Rhagoletis (Diptera: Tephritidae) are exemplars for sympatric speciation via host plant shifting. Flies in these species groups are morphologically similar and overlap in their geographic ranges, yet attack different, non‐overlapping sets of host plants. Ecological adaptations related to differences in host choice and preference have been shown to be important prezygotic barriers to gene flow between these taxa, as Rhagoletis flies mate on or near the fruit of their respective host plants. Non‐host‐related assortative mating is generally absent or present at low levels between these sympatrically diverging fly populations. 2. However, some Rhagoletis taxa occasionally migrate to ‘non‐natal’ plants that are the primary hosts of other, morphologically differentiated fly species in the genus. These observations raise the question of whether sexual isolation may reduce courtship and copulation between morphologically divergent species of Rhagoletis flies, contributing to their prezygotic isolation along with host‐specific mating. 3. Using reciprocal multiple‐choice mating trials, we measured sexual isolation among nine species pairs of morphologically differentiated Rhagoletis flies. Complete sexual isolation was observed in eight of the nine comparisons, while partial sexual isolation was observed in the remaining comparison. 4. We conclude that sexual isolation can be an effective prezygotic barrier to gene flow contributing to substantial reproductive isolation between many morphologically distinct Rhagoletis species, even in the absence of differential host plant choice and host‐associated mating.     

Comparative response to citrus foliage and citrus fruit odour by wild and mass-reared sterile Mediterranean fruit fly males of a genetic sexing strain

In greenhouse behavioural assessments conducted at Seibersdorf, Austria that allowed flies the choice of different types of potted host trees and plants, it was found that mature laboratory mass-reared Mediterranean fruit fly males, Ceratitis capitata (Wied), (Dipt. Tephritidae), strain Vienna-42 (temperature-sensitive lethal genetic sexing strain) showed the same preference for orange foliage, over other plants types and parts, as wild males. They aggregated in leks, released pheromone, and courted approaching females almost exclusively on orange foliage. Even though a number of other trees and plants were available to Vienna-42 males, less than 1% of these sexual activities occurred on the foliage or other plant parts of non-Citrus trees and not a single mating occurred there. In choice tests carried out in Chios, Greece in large field cages housing naturally planted orange trees, it was determined that mature and sterile Mediterranean fruit fly males of the same sexing strain responded to the odour emanating from the flavedo (due to superficial cuts made in the fruit peel containing the oil glands) of ripening sweet, sour oranges and lemons, in a similar intensity, order of fruit preference and time of day as did mature males of the local wild population. Males of both strains were frequently observed contacting with their mouth parts the oils exuding from the cuts made in the flavedo of the orange peel, or exhibiting homosexual activity on, or near the cuts. The results show that mass-reared, sterile males of the Vienna-42, genetic sexing strain behave in a comparable way to wild males in an important component of habitat location (odour of host foliage and host fruit) and hence are competitive with them in these respects. None of the standard international quality control tests for sterile Mediterranean fruit flies evaluates this important habitat location behaviour, which is a prerequisite for the successful encounter of the sexes in a sterile release programme.     

Infestation of fruit by conspecific and heterospecific females deters oviposition in two Tephritidae fruit fly species

Tephritidae fruit fly larvae develop entirely in the host chosen by the females. To improve the fitness of their progeny, females would benefit from rejecting previously exploited hosts. Anastrepha fraterculus and Ceratitis capitata are two species of fruit flies having similar nutritional requirements and overlapping in their distribution. Previous studies found that competition between the larvae of these species might reach high levels, suggesting that cross-recognition would be an adaptive trait. In this work, we tested the ability of A. fraterculus and C. capitata females to recognize and avoid fruits previously infested by both conspecific and heterospecific females. In laboratory behavioural arenas, females were presented with fruits that had been previously exposed to either conspecific or heterospecific females. Then, we conducted choice and non-choice assays to compare the response of A. fraterculus and C. capitata females to infested and non-infested fruits. In non-choice tests, the females from both species rejected fruits previously infested by conspecific and heterospecific individuals. However, the rejection occurred at different steps of the sequence leading to oviposition: A. fraterculus showed a lower rate of visits to infested fruits, whereas C. capitata visited both infested and non-infested fruits, but the latency to visit a fruit and the rejection frequency were higher and the duration of the visit to infested fruit was lower. In choice assays, the rejection of heterospecific infested fruit was higher than that of conspecific infested fruits, for both species. Our results suggest that, regardless of the sensory mechanism used by females, the recognition of previous infestation is bidirectional and females of both species, belonging to different genera, recognize fruit infested by heterospecifics. These responses indicate that cross-recognition, supposedly a highly beneficial trait, could be occurring in nature, thus reducing interspecific competition and contributing to the coexistence of these species.     

Whole body extract of Mediterranean fruit fly males elicits high attraction in virgin females

The search for effective female attractants emanating from the host or body of fruit flies has been an area of intensive research for over three decades. In the present study, bodies of male Mediterranean fruit flies, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), were extracted with diethyl ether or methanol and subjected to gas chromatography–mass spectrometry. Analysis revealed substantial qualitative and quantitative differences between males from a laboratory culture and wild males captured alive in an orchard. Most notably, the hydrocarbon sesquiterpene (±)‐α‐copaene, which is known to be involved in the sexual behaviour of the species, was found in substantial amounts in wild males, but was not detected in laboratory males. In laboratory tests, 15 laboratory or wild male equivalents of diethyl ether extracts or combined diethyl ether and methanol extracts, or, to a lesser extent, methanol extracts alone, were found to attract virgin females. In a citrus orchard, traps baited with combined diethyl ether and methanol extracts of wild males attracted significantly more virgin females than traps baited with various doses of pyranone or blends of other compounds identified in the extracts or reported in the literature, such as ethyl acetate, ethyl‐(E)‐3‐octenoate, and 1‐pyrroline. Traps baited with blends of compounds, however, displayed substantial attractiveness compared to control (non‐baited) traps. These results are important for better understanding the mating system of C. capitata as well as for further improving existing monitoring and control systems.