Host selection by a mycophagous fly and its impact on fly survival

Epichloë fungi interact with Botanophila flies in a co-evolved mutualism that appears to be analogous to angiosperm pollinator-parasite systems. Female flies act as vectors of fungal spores as they lay eggs on the fungi which grow within and on grass stems. To understand selective pressures operating on choice of host fungi by flies, we monitored the interaction of two species of Epichloë with Botanophila at two sites in southern England. At one site (Ashurst), the hosts Epichloë clarkii and Epichloë baconii co-occurred, while at a second site (Farm Gate), only E. clarkii was present. Flies ovipositing on E. baconii showed preferences for large, somewhat isolated fungi occurring within small clumps of grass. Flies ovipositing on E. clarkii at the Ashurst site did not select hosts based on any physical features of fungi that we measured. However, flies ovipositing on E. clarkii at the Farm Gate site tended to prefer large fungi that occurred within small clumps of grass. Fungal characteristics preferred by the flies showed no correlation with egg/larval survivorship. Insect survival on E. clarkii varied between sites and was related to differences in egg dispersion among fungi at the two sites. The absence of an alternate host ( E. baconii ) was associated with greater aggregation of eggs on E. clarkii at the Farm Gate site and these insects experienced higher levels of mortality at this compared to the other site.     

An analysis of the deterrent effect of aphids on cabbage root fly (Delia radicum) egg-laying

Abstract. 1. The cabbage root fly, Delia radicum (L.), was deterred from laying eggs on brassica plants with >250 cabbage aphid, Brevicoryne brassicae (L.), or peach-potato aphid, Myzuspersicae (Sulz.).
2. Flies did not lay on plants infested with >250 aphids.
3. Preparations of (E)-β-farnesene, the aphid alarm pheromone, deterred the flies from laying only at the extremely high dose of 32 mg/plant.
4. Although M. persicae secreted large (1 ng/insect) amounts of alarm pheromone and B. brassicae extremely small (<0.01 ng/insect) amounts, both aphids equally deterred D. radicum from laying.
5. The deterrent effect appeared to result from the aphids physically disturbing the flies during host-plant selection.     

Aggregated oviposition in the Simulium damnosum complex is mediated by eggs in a laboratory bioassay

Abstract. . The phenomenon of aggregated oviposition in blackflies (Diptera: Simuliidae) was investigated under controlled laboratory conditions, using wild-caught females of the Simulium damnosum complex in Sierra Leone. A method was developed for inducing Simulium females to lay eggs, and used as a bioassay to measure the responses of gravid S. damnosum s.l. to freshly laid eggs of the same species complex. In a series of two-choice tests, significantly more ovipositing flies chose substrates already containing eggs over control substrates ( P = 0.004). The time from introduction of flies into the oviposition system to the onset of egg-laying was significantly less when eggs were already present ( P = 0.049). Flies responded more quickly when more eggs were present and the relationship between egg-batch number and the time of this response was curvilinear ( P = 0.012). Ecological advantages and disadvantages of such aggregation behaviour and the possible role of semiochemicals in its mediation are discussed.     

Oviposition response of Simulium reptans (Diptera: Simuliidae) to the presence of conspecific eggs

Abstract.

• 1 The oviposition behaviour of Simulium reptans L. is described from two sites on the River Spey, Scotland. Female aggregations were observed immediately downstream of oviposition sites and were composed mainly of gravid flies (range 60–80%0) together with smaller numbers that were either freshly bloodfed (range 2–17%) or infected with mermithid nematodes (range 0–30%).
• 2 The time from landing on the oviposition sites to the onset of oviposition was recorded. The time in the presence of greater than 1-day-old eggs did not significantly differ from sites with no eggs present. However, the presence of freshly laid or I-day-old eggs significantly shortened the time to onset of oviposition. The cues that elicit oviposition are unknown but it is speculated that they may involve a pheromone.

     

Influence of background on host-plant selection and subsequent oviposition by the cabbage root fly (Delia radicum)

The behaviour of female cabbage root flies during host plant selection was studied in the laboratory using brassica plants growing in backgrounds of bare soil, clover, grass, peas and four non-living materials. Gravid females landed about twice as often on brassica plants growing in bare soil than on comparable plants growing amongst non-host plants. Once a receptive female landed on a brassica plant, the female made, on average, four ‘spiral flights’ and two jumps on and off the plant before laying alongside the plant. Surrounding a brassica plant with a diverse background altered the behaviour of the flies, so that the spiral flights around the host plant were replaced by short hops between nearby vertical objects. The loss of contact and recontact with the host plant then prevented the females from accumulating sufficient contacts with the host plant to be stimulated to lay. Spiral flights around host plants appear to determine whether or not flies will lay alongside host plants. Flies in mixed plantings have a reduced rate of settling on the host plant, and a higher rate of locomotion, because they land frequently on non-host plants. Hence, visual stimuli appeared to have greater effects than, chemical or mechanical barriers in deterring flies from laying alongside brassica plants in diverse backgrounds. In ‘choice’ situations, backgrounds of real plants reduced oviposition alongside brassica plants by at least 50%. In ‘no-choice’ situations, flies laid similar numbers of eggs alongside all brassica plants irrespective of plant background or plant size. If numbers of fly eggs are to be reduced on commercial brassica crops by undersowing the crops with clover, plants growing in bare soil may also have to be included to provide the flies with sites preferred for oviposition.     

The effects of the timing of exposure to cadmium on the oviposition behavior of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>

Animals can greatly increase their fitness by choosing oviposition sites free from damaging substances such as the heavy metal cadmium (Cd). Previous research has shown that Drosophila melanogaster reared on uncontaminated media lay fewer eggs on substrates containing Cd. In this study, we examined the effects of prior exposure to Cd on oviposition site preference. We tested flies that had been exposed to Cd continuously from egg to adult, only during pre-adult stages, or for the 4 days of adulthood prior to testing. We found that flies avoid laying eggs on Cd-contaminated medium and also that flies exposed to Cd as adults laid significantly fewer eggs than controls never exposed to Cd. We did not observe consistent changes in oviposition as a result of pre-adult exposure alone. Our results suggest that the presence of cadmium in the environment, even at low doses and for short periods of time during adulthood, can harm the fitness of Drosophila melanogaster.     

The oviposition sites of five leafhopper species (Horn. Auchenorrhyncha) on Holcus mollis and H.lanatus

Abstract. 1. A study was made of the oviposition sites of five leafhopper species Diplocolenus abdominalis, Recilia coronifera, Errastunus ocellaris, Elymana sulphurella (Cicadellidae) and Dicranotropic hamata (Delphacidae) on the grasses Holcus mollis and H.lanatus.
2. Experiments were made in the laboratory, each insect species being provided simultaneously with a complete plant of both Holcus species.
3. Distribution of eggs within the leaf sheath and leaf blade, and their height above ground-level, were recorded.
4. The presence and size of egg groups were noted, as well as the angle of eggs to the longitudinal axis of the leaf.
5. On the basis of comparison of the vertical distribution of the eggs with availability of plant tissues, the five species can be divided into three groups: (a) D.abdominalis and R.coronifera , (b) D.hamata and E.ocellaris , (c) E.sulphurella. Species within a group, i.e. with similar distributions of eggs, have other features separating their oviposition sites.     

Characteristics and distribution of opposition sites of the grasshopper Zonocerus variegatus (L.)

Abstract. 1. The dry-season population of Zonocerus in southern Nigeria oviposits in dense aggregations under single or clumped woody plants or on the edge of tall continuous stands of vegetation. No egg pods were found oviposited singly.
2. The vegetation under which oviposition occurs is the most common available at the time of the year. There seems to be no selection of particular species of plants so long as they provide good low shade.
3. Oviposition can take place in a wide range of gravelly soils.
4. Topography is not important as long as the oviposition site is on high ground away from damp areas subject to flooding.
5. There was a mean of 618 pods per oviposition site from forty-one sites which were dug up, and there was a significant correlation between the area of a site and the number of pods.
6. There was a marked tendency for oviposition sites to be found close to cassava. At two localities of approximately 20 ha each 90% of the sites were found within 30 m of cassava.
7. It is suggested that permanent shade is the most important factor in the selection of a site and this is probably related to the survival of the eggs during their 6–7 months development.     

Spatial contagion drives colonization and recruitment of frogflies on clutches of red-eyed treefrogs

Spatial contagion occurs when the perceived suitability of neighbouring habitat patches is not independent. As a result, organisms may colonize less-preferred patches near preferred patches and avoid preferred patches near non-preferred patches. Spatial contagion may thus alter colonization dynamics as well as the type and frequency of post-colonization interactions. Studies have only recently documented the phenomenon of spatial contagion and begun to examine its consequences for local recruitment. Here, we test for spatial contagion in the colonization of arboreal egg clutches of red-eyed treefrogs by a frogfly and examine the consequences of contagion for fly recruitment. In laboratory choice experiments, flies oviposit almost exclusively on clutches containing dead frog eggs. In nature, however, flies often colonize intact clutches without dead eggs. Consistent with predictions of contagion-induced oviposition, we found that flies more frequently colonize intact clutches near damaged clutches and rarely colonize intact clutches near other intact clutches. Moreover, contagion appears to benefit flies. Flies survived equally well and suffered less parasitism on clutches lacking dead eggs. This study demonstrates how reward contagion can influence colonization dynamics and suggests that colonization patterns caused by contagion may have important population- and community-level consequences.     

Phycomycetes parasitizing the ovaries of blackflies (simuliidae)

The developmental stages of the fungus in the ovary and freshwater are discussed. Based on the evidence to date the simuliid host is infected in the early larval instars. Parasitism results in a destruction of the ovarian tissues with no or few eggs produced. The fungus replaces the ovarian tissue and produces massive numbers of spores which are released by the female fly at the time of oviposition. After oviposition, infected flies do not seek a blood meal. Longevity of the host does not appear to be affected by the presence of the fungus. The spores released by the fly during oviposition did not withstand desiccation and did not develop on the artificial media tested; development occurs only in water. Development of sporangia takes 5 to 7 days at 4°C, and 2 to 3 days at temperatures above 10°C. The developmental process in spores from S. verecundum was followed from the formation of sporangia to the opening of discharge tubes which zoospores were presumed to be released but not observed.     

Field Studies of Entomophthora (Zygomycetes: Entomophthorales)—Induced Behavioral Fever in Musca domestica (Diptera: Muscidae) in Denmark

House flies were collected over 3 days (three to five times per day) from specific sites on a dairy farm with a range of high to low temperatures. Flies were held individually to determine whether the distribution of fungus-infected (Entomophthora muscae and E. schizophorae) house flies differed according to the stage of infection and temperature. All but 2 of 396 infected flies (99.5%) had E. muscae. More E. muscae-infected flies collected from cool areas were in later stages of infection (i.e., dying 0–2 days after capture), whereas flies collected on sun-exposed surfaces tended to be in earlier stages of infection (i.e., dying 6–8 days after capture). Most flies died 3–5 days after capture and were consequently in the middle stages of infection. A mark and release experiment was conducted to determine whether E. schizophorae-inoculated flies frequented surfaces with higher temperatures than did uninfected control flies. About 3000 yellow-marked house flies inoculated with E. schizophorae and 3000 blue-marked control flies were released in an enclosed swine farrowing barn. Significantly more inoculated flies were recorded on the heat lamps than flies in the control group. The results suggest that behavioral fever occurs in the field for flies infected with both E. muscae and E. schizophorae and that flies can cure themselves of infection through the use of artificial heat sources.     

Entomophthora infecting wheat bulb fly at Rothamsted, Hertfordshire, 1967–71

Entomophthora dipterigena, E. hylemyiae and, most commonly, E. muscae infected wheat bulb flies at Harpenden, Hertfordshire, between 1967 and 1971. The mean annual percentages of infected flies caught were 19, 0, 1, 29 and 16 respectively. These showed an increase with increasing host density. In 1970, two-thirds of the female flies were killed by E. muscae before they laid any eggs.
Conidiophores developed in most flies infected with E. muscae during the 2 weeks after peak emergence of flies. Subsequently, resting spores developed in a progressively increasing proportion of infected flies. Resting spores developed in only six of 130 infected males compared with 103 of 244 infected females in 1970 and 1971. The physiological age of infected flies probably determined whether conidiophores or resting spores developed.     

Abnormal egg-laying behaviour of female carrot flies (Psila rosae) induced by the fungus Entomophthora muscae

Female carrot flies (Psila rosae F.) infected with Entomophthora muscae (C.) Fres. on days 1 to 4 of adult life were unable to lay their eggs near the food plant. Flies infected on day 5 of adult life showed an egg-laying behaviour similar to that of uninfected flies. E. muscae-infected female carrot flies caught in the field showed an egg-laying behaviour comparable to that of the flies infected in the laboratory during the first day of their live.Eggs laid by infected females were equally as fertile as eggs laid by uninfected females, and developed into normal flies.It is concluded that female carrot flies infected with E. muscae in the field do not in general contribute to the development of the carrot fly population, due to their abnormal egg-laying behaviour.

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Zusammenfassung Wenn Weibchen der echten Möhrenfliegen (Psila rosae) zwischen den ersten und der vierten Tag in dem Laboratorium mit E. muscae infiziert worden waren, könnten sie ihre Eier an den Futterpflanze nicht ablegen. E. muscae-infizierte Weibchen, die ins Feld gefangen worden waren, zeigten auch ein abweichendes Eiablageverhalten. Ihr Behalten war mit dem Verhalten von Fliegen vergleichbar, die an ihrem ersten Lebenstag im Laboratorium infiziert worden waren.Von infizierten Weibchen gelegte Eier waren ebenso fruchtbar wie die von nicht infizierten Weibchen gelegten Eiern. Es wird konkludiert, dass E. muscae-infizierten Weibchen der echten Möhrenfliegen der Entwicklung der Population keinen Beitrag leisten.

     

Post-alighting responses of Mexican fruit flies (Dipt., Tephritidae) to different insecticides in paint on attractive spheres

Abstract: Two new, comparatively safe insecticides (spinosad and imidacloprid) were compared with dimethoate (each at 1.5% active ingredient) for behavioural and mortality effects on Mexican fruit flies, Anastrepha ludens . Insecticide was mixed with sugar (as a feeding stimulant) and yellow latex paint (as an extending agent) applied to the surface of fruit-mimicking biodegradable 7 cm spheres made of sugar, flour and glycerin. Flies feeding on spinosad-treated spheres did not differ from flies feeding on untreated spheres in post-feeding intra-tree flight capability, amount of oviposition or mortality. Flies that fed on imidacloprid- or dimethoate-treated spheres for as little as 30 s experienced both high reduction in oviposition and high mortality compared with flies that fed on untreated spheres, and the flies from imidacloprid-treated spheres also showed a much reduced intra-tree flight capability. If baited with attractive odour, biodegradable yellow spheres treated with a surface coating of imidacloprid in latex paint and sugar could have potential for suppressing Mexican fruit flies on host trees.     

Oviposition aggregation pheromone in the Simulium damnosum complex

Abstract. Communal oviposition by the Simulium damnosum complex of Afrotropical blackflies (Diptera: Simuliidae) was investigated under controlled laboratory conditions, using wild-caught flies in Sierra Leone. Volatile compounds emitted by Simulium eggs were trapped using a closed collection system, and their attractiveness to gravid flies was tested in a two-choice behavioural bioassay. Significantly more female blackflies oviposited on substrates baited with freshly laid eggs (100% chose the baited substrate), or with the volatiles collected from freshly laid eggs (85% chose the baited substrate), in preference to the relevant control substrates. Substrates baited with volatiles from 12-h-old eggs were not significantly more attractive than controls (only 31% chose the baited substrates; P = 0.33). Gas chromatographic analysis of the egg volatiles consistently showed two peaks emanating from fresh eggs, but significantly lower amounts from 12-h-old eggs ( P <0.05). A novel system for collecting the volatiles from this and other blackfly species, as they laid eggs on a substrate in flowing water, is described. Volatiles collected using this method showed identical gas chromatographic profiles to those of fresh eggs alone, indicating that the flies themselves produced no other volatile chemical signals during oviposition. Evidently communal oviposition by S. damnosum s.l . was mediated by a pheromone emanating from fresh eggs. The role of pheromone-mediated egg aggregation in blackfly ecology is discussed, and its possible manipulation is considered.     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

Avoidance of occupied hosts by the Hessian fly: oviposition behaviour and consequences for larval survival

Abstract 1. Unoccupied wheat plants and wheat plants occupied by conspecific eggs or larvae were presented to ovipositing female Hessian flies in choice tests.
2. The presence of conspecific eggs on the leaf surfaces of wheat plants did not appear to have any effect on the responses of ovipositing Hessian fly females.
3. The presence of conspecific larvae at the base and nodes of wheat plants for 1, 6, or 11 days had significant effects on Hessian fly oviposition. Eggs oviposited on plants were inversely proportional to larval densities and days of larval occupation.
4. Feeding by Hessian fly larvae is associated with several changes in wheat plants. One of these changes, the growth arrestment of the plant, was measured by recording the heights of plants used in oviposition tests. Plant heights were inversely proportional to both larval densities and days of occupation. Plant heights were directly proportional to eggs oviposited on plants.
5. The consequences of adult female avoidance of plants occupied by conspecific larvae were investigated by allowing females to oviposit on unoccupied plants and 1-day, 6-day, and 11-day larval occupied plants, then scoring at the end of the first larval instar the survival of the offspring that resulted from this oviposition.
6. Survival during the first larval instar was 88% for the offspring of females that oviposited on unoccupied plants, decreasing to 82, 31, and 4% on the 1-day, 6-day, and 11-day occupied plant treatments. On these four plant treatments, a positive correlation was found between larval performance (i.e. survival) and the preferences of ovipositing females.
7. On the four plant treatments, relationships between first-instar larval density and first-instar larval survival varied significantly. On unoccupied plants, survival was inversely proportional to density. On plants oviposited on at 6 days of larval occupation, survival was directly proportional to density.     

Avoidance of egg parasitism through submerged oviposition by tandem pairs in the water strider, Aquarius paludum insularis (Heteroptera: Gerridae)

Abstract.  1. Females of the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae) carry males on their backs and oviposit under water after copulation. This study focuses on the benefit  A. paludum insularis receives by ovipositing in tandem.
2. Males guarded females in tandem through to the end of oviposition in 85% of copulations. Females in tandem dived deeper than single females, and the density of A. paludum insularis eggs increased with water depth. The proportion of eggs parasitized by a scelionid wasp, Tiphodytes gerriphagus Marchal (Hymenoptera: Scelionidae) decreased with increasing water depth.
3. These results suggest that during oviposition guarding by males is beneficial for females, because it enables pairs to dive and lay eggs deeper and in oviposition sites where the risk of egg parasitism is lower.     

Age structure and abundance in populations of muscoid flies from a poultry facility in Southeast Brazil

Muscina stabulans, M. domestica, Chrysomya putoria, C. megacephala and Stomoxys calcitrans were the most abundant muscoid flies captured in a poultry facility in southeastern Brazil. We examined the gonadotrophic profiles of the females caught at different sites and different times and found that Mu. stabulans and M. domestica, the predominant species, presented similar gonadotrophic profiles only when captured on the manure under the cages, but very different and sometimes opposite gonadotrophic profiles when sampled from wooden posts, vegetation or electric cords. We also determined sex ratios and relative abundance for these two species and found significant differences between them. More than 50% of the females of both species of Chrysomya captured on manure carried eggs or exhibited signs of recent oviposition. The vast majority of S. calcitrans presented ovaries with eggs or signs of recent oviposition. A small proportion of them had ovaries in the recent emerged condition. Our data on ovarian stages, sex ratio and relative abundance allowed us to associate different gonadotrophic profiles with each site and characterize each site as a resting, ovipositing or mating site.     

Population dynamics of Siphoninus philyreae in California in the presence and absence of a parasitoid, Encarsia partenopea

Abstract. 1. Densities of the whitefly, Siphoninus phillyreae Haliday, remained at low levels at two sites where a parasitoid, Encarsia parknopea (Walker), was released on whitefly-infested ash, Fraxinus sp., and pomegranate, Punica granatum. Populations of S.phillyreae at control sites, where parasitoids were absent, were increasing in density at the beginning of the summer.
2. By mid-summer, E.partenopea had appeared at all four control sites, and densities of S.phillyreae declined to levels similar to those at the release sites.
3. In the absence of the parasitoid, the age structure of S.phillyreae populations was dominated by the egg stage, indicating a population increasing in density. After E.partenopea became abundant at a site, the age structure of the population had a decreasing proportion of young stages.
4. The changes in S.phillyreae age structures were explained by examining the percentage of fourth instar S.phillyreae from which E.partenopea , as opposed to adult whiteflies, emerged. After the parasitoid became abundant at a site, the majority of fourth instars produced parasitoids rather than whitefly adults. The result was that the adult S.phillyreae population declined, and few eggs were laid.
5. The dynamics of the interaction between E.partenopea and S.phillyreae were similar on ash and pomegranate.