Learning affects mate choice in female fruit flies

Learning in the context of mate choice can influence sexualselection and speciation. Relatively little work, however, hasbeen conducted on the role of learning in the context of matechoice, and this topic has been mostly ignored in insects eventhough insects have served as a prime model system in researchon sexual selection and incipient speciation. Extending recentwork indicating apparently adaptive learning in the contextof sexual behavior by male fruit flies (Drosophila melanogaster),I tested for the effect of learning on mate choice by femalefruit flies. Compared to young virgin females that experiencedcourtship by large males, young virgin females that experiencedcourtship by small males were more likely to mate with smalland large males in a test conducted a day after the experiencephase. These results, which are the first clear empirical demonstrationof learning in the context of mate choice by female insects,lay the foundation for research on the role of learning in insectsexual selection and speciation.     

Effects of larval diet on development rates and reproductive maturation of male and female Mediterranean fruit flies

Abstract Mediterranean fruit fly Ceratitis capitata (Diptera: Tephritidae) larvae are capable of developing in one of many hosts that may vary greatly in quality. We hypothesized that they will respond to the larval environment in a manner beneficial to their subsequent reproductive performance. Accordingly, we investigated the effects of various larval diets (varying in the amount of protein and sugar they contain) on the size, development time, nutritional status and reproductive maturation (ovarian development and onset of sexual behaviour) of females and males. We found that flies which undergo larval development in artificial host fruit that contain sugar and protein ('protein-fed') were larger, developed faster and emerged with more nutritional reserves than flies that were protein-deprived as larvae. Protein-fed males, regardless of their size, became sexually active before males that developed in hosts with no protein. Protein-fed females produced more mature eggs than protein-deprived ones. Moreover, protein-fed females tended to copulate sooner than females that developed in hosts with no protein. In addition, regardless of female larval diet, females with more mature eggs tended to copulate sooner than females with less mature eggs. In light of these results, the importance of the larval environment for adult reproductive success is discussed.     

Cuticular hydrocarbons from six species of tephritid fruit flies

Cuticular alkanes obtained from larvae and adults of six species of tephritid fruit flies, Anastrepha ludens, A. suspensa, Ceratitis capitata, C. rosa, Dacus cucurbitae, and D. dorsalis were analyzed by gas chromatography. The same four major alkanes were shown to be present by capillary GC-mass spectra in all Anastrepha and Ceratitis larvae. Profiles were obtained from individual larvae and adult specimens that showed statistical differences between species, whereas profiles of conspecific life forms including pupae of A. suspensa from different locations showed some similarities. Sexual dimorphism was not observed in alkanes extracted from adults. A novel series of alkadienes was found in all Anastrepha and Ceratitis but not in Dacus.     

Advancements in management of major fruit flies (Diptera: Tephritidae) in North Africa and future challenges: A review

Fruit flies are one of the most economically damaging pests of fleshy fruits worldwide. Two species of highest concern for fruit production in North Africa are the Mediterranean fruit fly (Medfly) Ceratitis capitata (Wied.) and the peach fruit fly (PFF) Bactrocera zonata (Saunders) (in Egypt and Libya only). Currently, both fruit fly species are mainly targeted by chemical applications of broad-spectrum contact insecticides. Despite the disparities in control efforts among North African countries, government and research are focused on reducing chemical reliance and adopt more environmentally friendly technologies. In this review, advances in integrated pest management (IPM) implementation against fruit flies are outlined for each country. In addition, challenges for future efforts are identified with emphasis on the efficacy of trapping for monitoring and control.     

Learning decreases heterospecific courtship and mating in fruit flies

Recent theory and data suggest that adaptive use of learning in the context of sexual behaviour could contribute to assortative mating. Experiments examining this issue indicated that male Drosophila persimilis that experienced courtship and rejection by heterospecific females exhibited significantly lower levels of heterospecific courtship and mating compared with those of inexperienced males. These results indicate that experience in the context of sexual behaviour in fruit flies could reduce gene flow between diverging populations, which may contribute to incipient speciation.     

Attractiveness of various colours to Australian tephritid fruit flies in the field

The response of tephritid fruit flies to variously coloured sticky traps was studied in the field in southeastern Queensland over three seasons (1978–81). Three species, Dacus tryoni (Froggatt), D. neohumeralis Hardy and D. cacuminatus (Hering), made up the majority of the flies captured. On a per unit area basis, a sticky flat trap was more efficient than a sticky wing trap and was adopted as the trap design. Daylight fluorescent (DF) Saturn Yellow captured significantly more flies than any of the other ten colours, or perspex and aluminium foil, tested, although the DF colours Lime, Blaze Orange and Emerald were also attractive. There was a significant correlation between the number of flies captured by a colour and the difference, in nanometres, between the peak reflected wavelength of the colour and 550 nm which is quoted as the peak wavelength reflected by green leaves. The three species did not differ in their responses to the coloured traps, and more males than females were captured. When either cuelure or methyl eugenol was added to the sticky traps no one colour emerged as significantly superior, but the yellow, and yellow/green DF colours consistently caught more flies. The attractiveness of coloured traps declined as the number of flies captured increased. The efficiency of Steiner and McPhail traps was not increased by the addition of any attractive colour. Of various possible trap shapes of equivalent surface area, circular and square traps captured more flies than did triangular, rectangular and diamond shaped traps. When spheres were tested, those of 10 cm diameter captured more flies than those of 5 cm diameter, and more flies were caught on black, than on yellow or green spheres.

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Résumé Plusieurs espèces de téphritidées, principalement Dacus tryoni, D. neohumeralis et D. cacuminatus, ont été capturées avec des pièges gluants colorés dans le S.E. Queensland (Australie) pendant 3 saisons (1978–81). Des pièges plats de 15×20 cm ont capturé plus de mouches/cm² que des pièges ailés formés par 2 plans à 90°. Les couleurs jaune-vert ont été préferées par les 3 espèces, le jaune saturne lumière du jour fluorescent ayant été le plus attractif. Une corrélation significative a été trouvée entre le nombre de mouches capturées par une couleur et la différence en nanomètres entre le mode des longueurs d'onde reflétées par cette couleur et 550 nm considéré généralement comme le mode des longueurs d'onde reflétées par les feuilles vertes. Les 3 espèces ont montré les mêmes réponses aux pièges colorés et le nombre de mâles avait tendance à excéder celui des femelles. Quand de la cuelure ou du méthyl eugénol étaient ajoutés aux pièges gluants, aucune couleur n'était significativement supérieure, mais les couleurs fluorescentes lumière du jour jaune et jaune-vert ont capturé nettement plus de mouches que le bleu, l'orange, le rouge, le noir, le blanc, la feuille d'aluminium ou le perspex clair. Le taux de capture diminuait plus le nombre de mouches sur le piège augmentait. Ceci peut expliquer la diminution apparente de la discrimination entre les couleurs quand des leurres mâles sont posés sur les pièges. Des expériences avec des pièges tachetés et un retrait quotidien des mouches ont montré que la diminution d'attractivité était une réponse à un stimulus visuel. Après plusieurs jours d'exposition, un piège de couleur relativement peu attractive, capturant des mouches à un taux faible mais appréciable, a capturé un nombre de mouches voisin de celui d'un piège de couleur attractive qui avait commencé par capturer un grand nombre de mouches, mais avait vu baisser son attractivité par suite de la présence des mouches captives. Les données hebdomadaires sur les captures n'ont pas présenté de différences suivant les couleurs, mais la sommation des captures quotidiennes (avec retrait quotidien des mouches) sur la même période a montré des différences d'attractivité significatives.Les pièges de Steiner et de McPhail n'ont présenté aucun taux de capture différent quand ils étaient peints sur les 2/3 inférieurs avec des couleurs attractives jaune-vert.Des sphères noires ont capturé plus de mouches que des sphères rouges, jaunes ou vertes, et des sphères de 10 cm de diamètre étaient plus attractives que celles de 5 cm.Quand un choix de pièges plats de la même couleur et de la même surface mais de formes diverses a été proposé aux mouches, les pièges circulaires et carrés ont capturé plus de mouches que les pièges triangulaires, rectangulaires ou rhombiques.

     

Effects of fish predation and seaweed competition on the survival and growth of corals

On Caribbean coral reefs, high rates of grazing by herbivorous fishes are thought to benefit corals because fishes consume competing seaweeds. We conducted field experiments in the Florida Keys, USA, to examine the effects of grazing fishes on coral/seaweed competition. Initially, fragments of Porites divaracata from an inshore habitat were transplanted into full-cage, half-cage, and no-cage treatments on a fore-reef. Within 48 h, 56% of the unprotected corals in half-cage and no-cage treatments (62 of 111) were completely consumed. Stoplight parrotfish (Sparisoma viride) were the major coral predators, with redband parrotfish (S. aurofrenatum) also commonly attacking this coral. Next, we transplanted fragments of P. porites collected from the fore-reef habitat where our caging experiments were being conducted into the three cage treatments, half in the presence of transplanted seaweeds, and half onto initially clean substrates. The corals were allowed to grow in these conditions, with concurrent development of competing seaweeds, for 14 weeks. Although seaweed cover and biomass were both significantly greater in the full-cage treatment, coral growth did not differ significantly between cage treatments even though corals placed with pre-planted seaweeds grew significantly less than corals placed on initially clean substrate. This surprising result occurred because parrotfishes not only grazed algae from accessible treatments, but also fed directly on our coral transplants. Parrotfish feeding scars were significantly more abundant on P. porites from the half and no-cage treatments than on corals in the full cages. On this Florida reef, direct fish predation on some coral species (P. divaracata) can exclude them from fore-reef areas, as has previously been shown for certain seaweeds and sponges. For other corals that live on the fore-reef (P. porites), the benefits of fishes removing seaweeds can be counterbalanced by the detrimental effects of fishes directly consuming corals. Received: 31 May 1997 / Accepted: 2 September 1997     

Evolution of delayed reproductive senescence in male fruit flies: Sperm competition

Populations ofDrosophila melanogaster that had been subjected to long-term selection favoring either delayed or rapid senescence were compared with respect to age-specific components of male reproductive success involving sperm competition. These components of reproductive success were divided into those related to sperm defense (protection of sperm from other males), and into those related to sperm offense (ability to mate with previously mated females and to displace the sperm of other males). Males were tested at four ages ranging from 1–2d to 5–6 wk after eclosion. Several aspects of sperm defense capability showed clear evidence of senescent decline. Furthermore, males from populations selected for delayed senescence were superior to males from control (rapid senescence) populations with regard to components of sperm defense. The superiority of males from populations with delayed senescence either increased as a function of male age, or was present at all ages tested. These results indicate that the rate of reproductive senescence in maleD. melanogaster can be altered in predictable directions by artificial selection. There were no differences between selection regimes with regard to sperm offense, and most components of sperm offense did not show clear evidence of senescence. The improved late-age reproductive success of males from populations selected for delayed senescence did not appear to entail any cost or trade-off at early ages with respect to the reproductive traits examined in these experiments.     

Ovipositional enhancement through socially facilitated behavior in Rhagoletis pomonella flies

Apple maggot females, Rhagoletis pomonella (Walsh), were maintained (with males) singly or in groups of 5 in laboratory cages without fruit from eclosion until 15 days of age (when males were removed). When hawthorn, apple or artificial fruit were introduced on Day 16, females caged in groups laid significantly more eggs per female, irrespective of fruit type, than females caged singly. A follow-up laboratory experiment revealed that the oviposition-enhancing effect of grouping was due largely or exclusively to caging regime on day of access to fruit and not to prior caging regime. Subsequent laboratory-cage tests indicated that none of the following factors contributed significantly to the oviposition-enhancing effect of grouping under laboratory cage conditions: odor of fruit punctures, odor of eggs, odor of marking pheromone, or presence of other females on fruit, all as potential stimuli eliciting female attraction to fruit; or presence of fruit punctures, eggs, or marking pheromone, all as potential oviposition-enhancing stimuli following alighting on a fruit. The only contributing factor of significance uncovered here was the enhanced propensity of an arriving female to bore into a fruit shortly after encounter with an occupying female engaged in ovipositional behavior. We consider such enhancement as suggestive of socially facilitated egglaying behavior, which we discuss.     

The genetic effects of competition in seaweed flies

In spite of abundant evidence that intra- and inter-specific competition occurs in natural communities, there is surprisingly little to suggest it is a major force promoting genetic change. This report assesses the genetic effects of competition in two species of seaweed fly, Coelopa frigida and C. pilipes. In laboratory cultures of C. frigida the relative survival of heterozygotes at the Adh locus, which was being used as a marker for the large αβ chromosomal inversion, was greater than that of homozygotes. In monocultures of C. frigida this competitive superiority was dependent on larval density. At low densities facilitation was seen, whereas at high larval densities there was competition. In mixed cultures of the two species, interspecific competition contributed to the differential mortality of C. frigida , and observations of natural populations suggested that competition may have similar effects to those described in laboratory culture. A possible mechanism involving the supply of nutritive microorganisms is proposed to underly both intra- and inter-specific competition. In seaweed flies, competition and the consequent differential mortality appear to be forces maintaining rather than reducing genetic variation.     

Effects of diet and activity on lipid levels of adult Mediterranean fruit flies

Abstract. The objectives of this study were to determine whether adult Mediterranean fruit flies, Ceratitis capitata Wiedeman (Diptera: Tephritidae), are capable of synthesizing lipids, and whether adult diet affects this ability.Lipid levels in females fed protein and carbohydrate or carbohydrate alone declined significantly from emergence to the fourth day of life and then rose back to teneral levels on the fifth day, before oviposition took place on the sixth day.In males fed protein and carbohydrate, lipid levels initially declined as males aged and then stabilized.In carbohydrate-fed males lipid levels declined following emergence and recovered somewhat by the sixth day.Lipid levels declined significantly when flies underwent post-emergence starvation, but after substantial feeding on the above-mentioned diets they eventually (within 6–7 days) reached teneral levels in all experimental groups.Multivariate analysis of variance revealed that differences in lipid contents are primarily related to the flies' age, which corresponds to the various sex-specific activities these flies exhibit.Average lipid investment in eggs was found to equal teneral lipid levels in females.Without lipogenic abilities, oviposition would completely deplete female lipid reserves.We conclude that adult medflies are capaple of lipid synthesis, and that this capability is modulated by individual and sex-specific activity patterns.     

The effect of social facilitation on foraging success in vultures: a modelling study

The status of many Gyps vulture populations are of acute conservation concern as several show marked and rapid decline. Vultures rely heavily on cues from conspecifics to locate carcasses via local enhancement. A simulation model is developed to explore the roles vulture and carcass densities play in this system, where information transfer plays a key role in locating food. We find a sigmoid relationship describing the probability of vultures finding food as a function of vulture density in the habitat. This relationship suggests a threshold density below which the foraging efficiency of the vulture population will drop rapidly towards zero. Management strategies should closely study this foraging system in order to maintain effective foraging densities.     

Biological attributes of three introduced parasitoids as natural enemies of fruit flies,genus Anastrepha (Diptera: Tephritidae)

The biological attributes of three introduced species of parasitoids which attack the fruit fly Anastrepha ludens were evaluated. Larvae and eggs of A. ludens were exposed to larval parasitoids Diachasmimorpha longicaudata and D. tryoni and the egg parasitoid Fopius arisanus. Parasitoid longevity and fecundity were determined using larvae and eggs of A. ludens. Likewise, the parasitism rates of these parasitoid species in infested host fruits were recorded. The intrinsic rate of increase for F. arisanus was 0.1019 followed by D. tryoni with a rate of 0.1641 and D. longicaudata with the highest rate of 0.2233. Although F. arisanus females had the highest levels of fecundity, only 50% of them remained alive until reproductive age. These results in combination with the longer generation time (in comparison with D. longicaudata and D. tryoni), can be considered as the most important factors explaining F. arisanus reduced rate of increase. However, we note that oviposition activity caused egg mortality which reduced Anastrepha egg hatch by ca. 20%. This result suggests that F. arisanus has a high potential as a natural enemy of A. ludens, in accordance with our research efforts to develop a new F. arisanus strain specialized for development in Anastrepha eggs. The results show that D. tryoni is not a good candidate for biological control of Anastrepha. Meanwhile, D. longicaudata continues to be the most important exotic parasitoid for suppression of Anastrepha fruit fly populations.     

Use of fruit wounds in oviposition by Mediterranean fruit flies

Casual observations suggested that female Mediterranean fruit flies (Ceratitis capitata Wiedemann) exploit fruit wounds (including pre-existing oviposition punctures) as oviposition sites. This behaviour was quantified under field conditions in a citrus grove on the Greek island of Chios. Fruit wounds influenced oviposition behaviour in three ways. First, females were more likely to land on oranges (Citrus sinensis) that were artificially wounded than into unwounded control oranges. Second, having landed, females were more likely to attempt oviposition into a wounded orange than into control oranges. Third, females that attempted oviposition into wounded oranges usually did so directly into or very near the wound. The diameter and depth of the wound significantly influenced the tendency for female flies to land on a fruit but not their propensity, having landed, to attempt oviposition in or near the wound. The significance of this behavior in nature and implications for management of the Mediterranean fruit fly are discussed.

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Résumé Des observations fortuites nous ont suggéré que C. capitata Weidemann utilise les blessures des fruits, y compris d'anciennes piqûres de ponte, pour déposer ses oeufs. Ce comportement a été quantifié dans une orangeraie de l'île grecque de Chios. Les blessures des fruits influent sur la ponte de 3 façons. D'abord, les femelles atterissent plus fréquemment sur les oranges, Citrus sinensis, artificiellement blessées que sur les fruits intacts. Ensuite, après atterrissage, les femelles tentent plus souvent de pondre sur une orange blessée que sur des oranges intactes. Enfin, les femelles pondent directement dans la blessure ou à proximité. Le diamètre et la profondeur de la blessure influent sur la tendance à l'atterrissage, mais non sur les tentatives de ponte dans la blessure ou près d'elle. La signification de ce comportement dans la nature et ses conséquences sur la lutte contre C. capitata sont discutées.

     

Hosts of Mediterranean fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) from Broome and the Broome Peninsula, Western Australia

Abstract  In the Kimberley region of Western Australia, Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) is only established at Broome. From 1998 to 2001 extensive fruit sampling was carried out at Broome during a trial investigating the eradication of medfly from the Kimberley using the sterile insect technique. Fruit was collected from 82 native and introduced plants growing in and surrounding the town of Broome and held for emergence of medfly. Eighteen plants, including the native tree Terminalia petiolaris , were found to be medfly hosts . On the basis of abundance, fruiting phenology and host suitability, eight species (kumquat Fortunella japonica , mango Magnifera indica , Barbados cherry Malpighia glabra , orange jessamine Murraya paniculata , guava Psidium guajava , Pacific almond Terminalia catapa , blackberry tree Terminalia petiolaris and yellow oleander Thevetia peruviana ) were the most important hosts ensuring medfly survival and population growth. Despite medfly being reared from T. petiolaris , it is likely that medfly can only maintain populations in areas close to human habitation, and eradication from these areas would lead to eradication from the whole Kimberley region.     

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.