The culture ofEntomophthora muscae (C) Fres. in carrot flies (Psila rosae F.) and the effect of temperature on the pathology of the fungus

A method for maintaining anin vivo culture ofEntomophthora muscae (C) Fres. on its original host, adult carrot flies (Psila rosae F.), is described. The lethal time for adult carrot flies was greatly influenced by temperature, both for infected and for uninfected flies. In the range 8.2°C–20.2°C the LT₅₀ for infected flies was about 5.4 times shorter than the estimated average life-span for uninfected flies. The discharge of primary spores was also strongly dependent on temperature. The total number of primary spores discharged per fly at 100% RH and in darkness ranged between 1.2×10⁴ and 9.6×10⁴ with a mean of 5.1×10⁴.      

Relationship between root volatiles of some carrot cultivars and their resistance to the carrot fly, Psila rosae

Field experiments investigated the resistance of some carrot cultivars to Psila rosae. In addition, headspace vapour and steam distillate from the roots of resistant and susceptible varieties were compared by gas-liquid chromatography. The field data confirmed that resistance may operate by decreasing the numbers of eggs laid indicating a nonpreference by the female Psila. Root resistance to the larva was also confirmed but the mechanism was unclear. A new finding was that root resistance is independent of the effect of egg laying, some cultivars evincing one or the other effect and some such as Regulus Imperial displaying both. It was clear that root resistance to the larva is the crucial prerequisite in breeding resistant varieties.One consistent difference was detected by the chemical comparisons: intact roots of resistant varieties released substantially less volatiles. Specifically, Regulus released almost five times less of the volatiles already shown to positively influence host-finding behaviour by the larva.

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Résumé La résistance à P. rosae de quelques cultivars de carotte a été étudiée en plein champ. Parallèlement, les substances volatiles diffusées et celles extraites par la vapeur des racines de variétés résistantes et sensibles, ont été comparées en chromatographie gaz-liquide (GLC). Les résultats en champ ont confirmé que la résistance peut être due à une diminution du nombre d'oeufs pondus, révélant une absence d'attractivité pour les femelles de P. rosae. La résistance des racines aux larves a été aussi confirmée, mais les raisons n'en étaient pas claires. Un aspect nouveau est que la résistance des racines est indépendante de l'effet de la ponte, quelques cultivars présentant l'un ou l'autreeffet et certains, comme Regulus Imperial, manifestant les deux. Il est net que la résistance racinaire aux larves est la condition essentielle pour la sélection de variétés résistantes. Une différence importante a été mise en évidence par les comparaisons chimiques: les racines intactes de variétés résistantes libèrent nettement moins de substances volatiles. Précisément, Regulus a libéré 5 fois moins de substances volatiles déjà connues comme influençant positivement le comportement de découverte de l'hôte par la larve.

     

Effect of the entomopathogenic fungus,Entomophthora muscae (Zygomycetes: Entomophthoraceae), on sex pheromone and other cuticular hydrocarbons of the house fly,Musca domestica

House fly (Musca domestica) males are highly attracted to dead female flies infected with the entomopathogenic fungus Entomophthora muscae. Because males orient to the larger abdomen of infected flies, both visual and chemical cues may be responsible for the heightened attraction to infected flies. Our behavioral assays demonstrated that the attraction is sex-specific-males were attracted more to infected females than to infected males, regardless of cadaver size. We examined the effect of E. muscae on the main component of the house fly sex pheromone, (Z)-9-tricosene, and other cuticular hydrocarbons including n-tricosane, n-pentacosane, (Z)-9-heptacosene, and total hydrocarbons of young (7 days old) and old (18 days old) virgin females. Young E. muscae-infected female flies accumulated significantly less sex pheromone and other hydrocarbons on their cuticular surface than uninfected females, whereas the cuticular hydrocarbons of older flies were unaffected by fungus infection. These results suggest that chemical cues other than (Z)-9-tricosene, visual cues other than abdomen size, or a combination of both sets of cues might be responsible for attraction of house fly males to E. muscae-infected females.     

Entomophthora ferdinandii (Zygomycetes: Entomophthorales) Causing Natural Infections of Musca domestica (Diptera: Muscidae) in Argentina

The identity and activity of an entomopathogenic fungus belonging to the Entomophthora muscae species complex and infecting Musca domestica in poultry houses from La Plata, Argentina, is reported. Entomophthora caused natural infections between September 2001 and September 2003. Primary conidia of this fungus were on average 29.5 ± 1.2 × 23.4 ± 2.4 μm and contained, on average, 10.5 ± 0.1 nuclei (range: 7–15) with an average diameter of 4.8 ± 0.1 μm. This fungus is identified as E. ferdinandii Keller (this specific epithet includes a nomenclaturally required spelling correction); this is a first record of E. ferdinandii in South America and of any member of the E. muscae species complex from flies in Argentina.     

Observations on egg-laying behaviour of the banana weevil,Cosmopolites sordidus (Germar)

Banana weevil females laid on an average 2.7 eggs/week in rhizome material and 0.7 eggs/week in pseudostem material in the laboratory. At extremely high weevil population densities the egg-laying activity declined. Under controlled field conditions 0.7 eggs/week were laid in banana suckers and 1.3 eggs/week in stumps of harvested suckers. 25% of the weevil stages found in suckers in the field were eggs of which 78% were laid in the rhizome and 22% in the pseudostem base. The majority of eggs was deposited in the crown area of the rhizome followed by the remaining surface area of the rhizome, the walls of abandoned larval tunnels in rhizome and pseudostem and the leaf sheaths. 58% of the eggs found were considered accessible to egg predators.     

Production of haploid-haploid chimeras by sperm injection in the sawfly,Athalia rosae (Hymenoptera)

Mature unfertilized eggs (oocytes) dissected from the ovary of the sawfly Athalia rosae (Hymenoptera) begin parthenogenetic development if exposed to distilled water and produce haploid males. Injection of sperm into mature oocytes through the anterior pole resulted in karyogamy in a fraction of cases which developed as diploid females. No haploid-haploid chimeras due to independent participation of the injected sperm in development were produced. When sperm were injected through the posterior pole, however, fertilization never occurred but haploid-haploid chimeras were produced in a smaller fraction of cases. Both egg nucleus-derived and injected sperm-derived nuclei contributed in forming the germ cells of the chimeric males.     

Experiments inducing prospective polar body nuclei to participate in embryogenesis of the sawfly Athalia rosae (Hymenoptera)

Summary Mature eggs dissected from ovaries of unmated females of Athalia rosae (Hymenoptera: Tenthredinidae), if placed on a filter-paper soaked with distilled water, are activated and develop to haploid males. Occasionally, however, diploid females develop from these artificially activated eggs. Treatment of mature unfertilized eggs dissected from diploid females with ice-cold temperatures immediately before activation and with a high temperature (36° C) upon and immediately after activation resulted in the production of diploid males, diploid females, triploid females and gynandromorphs at high frequency. The same treatment of mature unfertilized eggs dissected from triploid females resulted in the production of only triploid survivors. These results, together with the results on the segregation of a marker mutation, yellow fatbody (yfb), appear to indicate that meiotic divisions were complete in the treated eggs, and that all four nuclei became potentially capable of participating in development with or without automictic fusion.Studies on the sawfly, Athalia rosae (Insecta, Hymenoptera, Tenthredinidae), part V     

Logistic growth of the host‐specific obligate insect pathogenic fungus Entomophthora muscae in house flies (Musca domestica)

Insect pathogenic fungi (IPF) need to overcome the host immune system in order to sporulate and ensure transmission to new hosts. Some IPF produce immunosuppressive toxins, whereas others rely on rapid fungal proliferation to kill the host by sheer fungal mass, resulting in a trade‐off between allocating resources to toxin production and fungal proliferation. The obligate entomopathogenic fungus, Entomophthora muscae sensu stricto, is host specific to the common house fly, Musca domestica. E. muscae grows as protoplast cells without cell walls and is not known to produce toxins. Here, we assessed the growth of E. muscae, in vivo, using real‐time PCR to measure the amount of a single‐copy actin gene. We find that E. muscae exhibits S‐shaped logistic growth between time post‐exposure and the number of fungal nuclei. The results show that E. muscae initially grows exponentially inside the host until depletion of available nutrient sources signifies the ‘limiting capacity’ where after the host is killed. This growth pattern differs markedly from toxin‐producing IPF species of Metarhizium and Beauveria in which maximal (plateau) growth and sporulation do not occur until well after the death of the host.     

Female-derived sex pheromone mediates courtship behaviour in the parasitoid Lariophagus distinguendus

Brassiceye^®traps baited with ethylisothiocyanate were modified and used to collect live adults of Delia radicum(L.) and Delia floralis(Fallén) (Diptera: Anthomyiidae) from the field to observe the prevalence of Entomophthora muscae(Cohn) Fresenius and Strongwellsea castransBatko & Weiser. The traps were highly effective and selective for D. radicumand D. floralis. Of the flies identified, 98.4% in 1996 and 93.7% in 1997 were either D. radicumor D. floralis. In 1997 the maximum mean catch was as high as 82 flies per trap per day, and more than 80% of these were females.During both seasons E. muscaecaused relatively high levels of mortality in adult populations of D. radicumand D. floralis. The fungus caused a total infection level of 17.9% in 1996 and 47.7% in 1997 with infection peaks of 82.4% in 1996 and 87.5% in 1997. Both years, a significant positive correlation was found between E. muscaeprevalence and temperature. One infection peak was observed for S. castransin 1996, and during that season the total S. castransinfection level was 18.0%. In 1997, the total S. castransinfection level was as low as 8.1%. There is no strong indication that the prevalence of E. muscaeor S. castransdiffers between either the fly species or sexes within species.     

The effect of fungus infection ofMetopolophium dirhodum [Hom.: Aphididae] on the oviposition behaviour of the aphid parasitoidAphidius rhopalosiphi [Hym.: Aphidiidae]

The oviposition behaviour of the hymenopterous parasitoidAphidius rhopalosiphi onMetopolophium dirhodum nymphs at successive stages of infection with the aphid-pathogenic fungusErynia neoaphidis was compared with that on uninfected nymphs. The frequency with which the parasitoid attempted to oviposit diminished in aphids inoculated with the fungus 3 days previously, that is within only 24 h of dying of the infection, but not in those infected only 1 or 2 days previously. The parasitoid did not attempt to oviposit in aphids killed by the fungus.      

Entomophthora leyteensis Villacarlos & Keller sp. nov. (Entomophthorales: Zygomycetes) infecting Tetraleurodes acaciae (Quaintance) (Insecta,Hemiptera: Aleyrodidae), a recently introduced whitefly on Gliricidia sepium (Jaq.) Walp. (Fabaceae) in the Philippines

Entomophthora leyteensis Villacarlos & Keller sp. nov., a species of Entomophthorales infecting the whitefly Tetraleurodes acaciae on Gliricidia sepium in the Philippines is described. Disease prevalence monitored weekly for 8 weeks indicated that the fungus could cause 8-31% infection within the whitefly population. Epizootics due to this fungus occurred in Inopacan, Leyte. Sampling live whitefly adults and dissecting them on glass slides for microscopic examination of fungal structures was found to give a better measure of prevalence than actual counts of infected insect cadavers. E. leyteensis is an important mortality factor for T. acaciae. Some speculations on the origin of the fungus are discussed here.     

Entomophthora apiculata (Thaxter) Gustafs. (Zygomycetes,Entomophthorales) as a pathogen of calypterate flies in Nigeria

Entomophthora apiculata (Thaxter) Gustafs. was isolated from the following naturally-infected dipterans: Musca domestica L. (s.l.) (Muscidae); Hemipyrellia fernandica Macquart and Chrysomya chloropyga f. putoria Weidemann (Calliphoridae). This fungus produced rhizoids in vivo but not in vitro. Although it was readily cultured in vitro, experimental infection could not be achieved consistently — which calls for intensive research into the biology of this and other Entomophthora species.     

Egg Deposition Behavior in the Haplodiploid Sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: Symphyta: Tenthredinidae)

The egg deposition behavior of the turnip sawfly, Athalia rosae (Hymenoptera: Symphyta), is described. Both unmated and mated females lay eggs individually inside of fresh young leaves of cruciferous plants. During an oviposition event, females exhibit a distinct pause in abdominal contractions just before the actual egg deposition act. Unmated females show a longer pause (11.31 s on average) than mated females (4.38 s on overall average). By employing an eye color mutation, the sex of the eggs laid by females was ascertained. Females mated once lay mostly fertilized (diploid female) eggs initially but begin to lay a considerable number of unfertilized (haploid male) eggs later in life. The laying of an unfertilized egg is associated with a longer pause (6.98 s on average) than the laying of a fertilized egg (3.76 s on average). These results are in contrast to previous reports on apocritan Hymenoptera, where the presence of a pause or a longer pause during oviposition was associated with the deposition of fertilized eggs rather than unfertilized eggs. The possibility that mated Athalia rosae females control fertilization and its implications for sex allocation strategies are discussed.     

The influence of natural enemies on brood production inIps typographus (Col. scolytidae) with special reference to egg-laying and predation byThanasimus formicarius (Col.: Cleridae)

The predatorThanasimus formicarius (L.) (Coleoptera, Cleridae) and its preyIps typographus (L.) (Coleoptera, Scolytidae) were studied in the laboratory and the field. In the laboratory, 11T. formicarius laid 71–132 eggs (mean=162) during 66–123 days. During this time they ate 66–132I. typographus adults per pair (male +female). The number of eggs laid per female was not correlated with life span or the number ofIps eaten. In the field, predation byT. formicarius larvae and other natural enemies onI. typographus brood was studied in the last year of an outbreak. Caged and uncaged spruce bolts attacked byI. typographus were used, and pairs ofT. formicarius were released in the cages. The treatments were: uncaged bolts, caged bolts withoutT. formicarius, caged bolts with 4T. formicarius pairs, and caged bolts with 8T. formicarius pairs. The productivity ofI. typographus was highest in the caged bolts withoutT. formicarius (mean=4.5 offspring/female) and lowest in the uncaged bolts (mean=0.9 offspring/female). The density ofI. typographus galleries was similar in the different treatments. Hence, the variation in productivity between treatments could not have been due to differences in the levels of intraspecific competition. There was no difference in bark beetle productivity or density ofT. formicarius larvae between bolts with 4 pairs ofT. formicarius and bolts with 8 pairs (mean=2.5 offspring/female). This indicates that some kind of interference occurred betweenT. formicarius individuals (e.g. cannibalism) and that a maximum level of predation was reached. Predation by larvae ofMedetera spp. (Diptera, Dolichopodidae),Thanasimus spp. and other beetles, and parasitism by wasps (Hymenoptera, Pteromalidae) probably caused the low productivity in the uncaged bolts.      

Marking of host fruit by maleRhagoletis boycei Cresson flies (Diptera: Tephritidae) and its effect on egg-laying

MaleRhagoletis boycei flies (Cresson) have been shown previously to guard egg-laying punctures on host walnut (Juglans major) fruit. During their tenure,R. boycei males were observed to dip the posterior portion of their abdomen repeatedly toward the fruit surface. Closeup video analysis revealed that, during this dipping behavior, a clear viscous substance emanating from the cercus was deposited directly on the fruit. Laboratory assays indicated that the rate of abdomen-dipping was significantly higher on fruit bearing simulated egg-laying punctures than on control fruit and significantly higher in the area of the puncture than elsewhere on the fruit. In a final experiment, the occurrence of abdomen-dipping in the vicinity of a puncture was manipulated independently on separate punctures on the same fruit. On such fruit and in the absence of males, previously mated, reproductively mature females attempted oviposition significantly more often in or near a puncture exposed to males than in or near an unexposed control puncture. Possible functions of the putative male-marking behavior from the perspectives of both male and female are discussed.     

Independent evolution of the same set of characters in fig flies (Lissocephala,Drosophilidae)

Summary Comparison of species of African fig flies (Lissocephala, Drosophilidae) shows that an unusual phenotype has evolved independently when a particular environment prevails. Each time the adaptation is observed, associated morphological and behavioural characters also appear. Such convergence suggests that selection operates on sets of predetermined morphogenetic pathways.     

Control of Muscidae on cattle by flucythrinate ear-tags, the behaviour of these flies on cattle and the effects on fly-dislodging behaviour

In two trials using diary heifers, in the south of England, ear-tags, containing flucythrinate, completely controlled Haematobia irritans (L). Musca autumnalis De Geer on the face were reduced by 41% using one tag and by 76% using two tags per heifer. A single ear-tag reduced numbers of Hydrotaea irritans (Fallén) on the body by 90% and on the teats by more than 55%. Other minor pestering flies were similarly controlled. Control was sustained over the entire 14 weeks duration of the trials which effectively encompassed the fly season of the three main pests.Ear-flicks by the heifers were related to dislodging M. autumnalis from the face, and tail flicks to dislodging Hy. irritans from the belly. These are the two commonest species of nuisance flies and are known to carry pathogenic bacteria.A management plan intended to maximise effective use of insecticidal ear-tags while minimising risks of insecticide resistance is described.

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Résumé Des boucles, contenant du flucythrinate, ont protégé totalement des génisses pour la production de lait contre Haematobia irritans L. lors de deux traitements expérimentaux dans le sud de l'Angleterre.Les effectifs de Musca autumnalis De Geer sur la face des génisses ont été réduits de 41% en utilisant un boucles et de 76% en utilisant deux. Un seul boucles réduit les effectifs d'Hydrotaea irritans Fallén de 90% sur le corps et de plus de 55% sur les pis. Les effectifs d'autres mouches nuisibles de moindre intérêt ont été réduits eux aussi. L'efficacité a été maintenue pendant les 14 semaines du traitement ce qui déborde efficacement le période de vol des 3 principales espèces de mouches nuisibles. Les battements d'oreille des génisses sont destinés à chasser M. autumnalis du visage, et les coups de queue à chasser H. irritans du ventre.Ce sont les deux principales espèces de mouches nuisibles connues comme vecteurs de bactéries pathogènes.Un programme est proposé pour optimaliser l'efficacité des boucles, tout en limitant les risques de résistance aux insecticides.

     

Effect of prey density on behaviour and development of the predatory mosquito,Toxorhynchites towadensis

The effect of prey density on feeding behaviour, killing behaviour, and development of the predatory mosquito,Toxorhynchites towadensis, was investigated in the laboratory. The number of prey consumed per larva increased toward an upper asymptote as prey density increased. Prey consumption curves during fourth instar were concave at low prey densities but convex at high prey densities. This phenomenon was not observed during other instars. Killing without consuming any part of prey occurred at prey densities of 20 per container and over. The number of prey killed but not consumed increased linearly with the number of unconsumed prey in the container. Prey acquisition behaviour was not affected by prey densities during the prepupal period. Developmental time from first instar to adult emergence decreased with increasing prey densities, but remained constant at densities of 10 per container and over. Adult size increased with increasing prey densities but there was no effect at prey densities of 20 and over.