Activity and distribution of the mushroom phorid fly,Megaselia halterata,in and around commercial mushroom farms

The mushroom phorid fly, Megaselia halterata (Wood) (Diptera: Phoridae), is a key pest in mushroom farming in most parts of the world. Studies on the mushroom phorid fly have focused on its life history within mushroom growing houses, but little is known about the fly's activity outside mushroom growing houses. In this study, daily activity and distribution of adult M. halterata in the areas surrounding mushroom growing houses was studied using yellow sticky traps. Results suggest that M. halterata focuses its flight activity over turf areas rather than windbreaks and spent compost piles, possibly for mating purposes. Our study found no evidence of M. halterata breeding in turf areas surrounding mushroom growing houses. In addition, flight activity is highest in the afternoon until midnight at higher temperatures, yet at lower temperatures activity ceases after sunset. Establishing temperature and daylight thresholds for M. halterata flight activity may be useful in developing integrated pest management (IPM) tactics for this species. The most successful IPM tool that mushroom growers use at present is fly exclusion. Exclusion can be improved by focusing farm operations around temperature and daylight thresholds when fly activity is at its lowest.     

Bioassay for evaluating attraction of the phorid fly,Megaselia halterata to compost colonized by the commercial mushroom,Agaricus bisporus and to 1-octen-3-ol and 3-octanone

A bioassay chamber was developed to evaluate the attraction of the phorid fly pest,Megaselia halterata (Wood) (Diptera, Phoridae), to volatiles from mushroom compost colonized by the commercial mushroom,Agaricus bisporus (Fungi, Agaricaceae) and to its volatile constituents, 1-octen-3-ol and 3-octanone. The bioassay chamber consisted of a large round container which was attached, via glass tubes, to three removable sample cups situated below the container. Once a fly had entered a sample cup, the flatly cut glass tubes prevented the flies from readily returning to the release chamber and, in effect, was a pit fall trap which kept the flies in the cups until the flies were counted. Holes in the bottom of the cups and removal of 20 ml of air from the chamber to fill the entrance tubes with volatiles from the cups were necessary for the success of the bioassay. Mated and unmated femaleM. halterata and, to a lesser extent, mated males preferred spawned compost (compost well colonized by fungal mycelia) relative to unspawned compost or blank cups. Unmated males were not attracted to spawned compost. Males and females were not attracted to unspawned compost. No attraction of male or femaleM. halterata to the spawned compost volatiles, 1-octen-3-ol and 3-octanone, could be demonstrated under various experimental conditions. These two compounds seemed to deter females at high concentrations.     

Evaluation of the behavioural response of the flies Megaselia halterata and Lycoriella castanescens to different mushroom cultivation materials

A static‐air olfactometer was used to investigate the behavioural responses of adult female phorid [Megaselia halterata (Wood) (Diptera: Phoridae)] and sciarid [Lycoriella castanescens (Lengersdorf) (Diptera: Sciaridae)] flies to the commercial white mushroom, Agaricus bisporus (Lange) Imbach, grown on a standard pasteurised composted substrate. The attraction of the flies was measured in relation to four test materials: composted substrate spawned with A. bisporus mycelium for 4 days and 14 days, uncolonised composted substrate, and A. bisporus sporophores. The experiment was done according to a 4 × 4 × 4 Latin cube design, and the results were analysed using a generalised linear model. It was found that both the occasion on which a bioassay was run and the position of the olfactometer within a 4 × 4 array could affect the proportion of the fly population responding to a test material. Megalesia halterata preferred spawned compost to unspawned compost, and the level of response to compost spawned for 14 days was greater than to compost spawned for 4 days. In contrast, L. castanescens were attracted equally to all of the materials tested. Overall, L. castanescens showed a greater level of activity than M. halterata, and was more likely to enter the pitfall traps in the olfactometer. For both M. halterata and L. castanescens, the type of test material affected the numbers of adult flies of the F₁ generation that emerged from it following oviposition. The highest numbers of emerging M. halterata were obtained from a composted substrate spawned for 4 days, and none emerged from the unspawned compost. Emergence of L. castanescens was highest from the uncolonised composted substrate, and there was a negative relationship between emergence and the amount of mycelium in the composted substrate. The results are consistent with the use of volatiles in the detection of oviposition sites by both species; however, further studies of the materials will be necessary to determine precisely which oviposition cues the insects use.     

Development and reproductive capacity of the predatory mite <Emphasis Type="Italic">Parasitus consanguineus</Emphasis> (Acari: Parasitidae) reared on the larval stages of <Emphasis Type="Italic">Megaselia halterata</Emphasis> and <Emphasis Type="Italic">Lycoriella ingenua</Emphasis>

Development and reproduction of the predatory mite Parasitus consanguineus Oudemans et Voigts (Acari: Parasitidae) reared on a diet of first and second instars of Megaselia halterata (Diptera: Phoridae) or Lycoriella ingenua (Diptera: Sciaridae) were studied. Mites were allowed to feed on these diets until death. The developmental time of immature stages of P. consanguineus was significantly longer when reared on L. ingenua than on M. halterata larvae (8.3 vs. 7.9 days, respectively). Survival to adulthood of P. consanguineus reared on L. ingenua or M. halterata larvae was 63 and 49%, and mite fecundity was 17.8 and 12.3 eggs/female, respectively. Adult females reared on L. ingenua lived on average 6.9 days, whereas those reared on M. halterata lived for 5.7 days. Mite survival, female longevity and fecundity were significantly different among the two diet types.     

Substrate dependent larval development and emergence of the mushroom pests Lycoriella auripila and Megaselia halterata

Pasteurized, spawned, full-grown and immediately-cased full-grown compost were simultaneously exposed to natural populations of the mushroom pests Lycoriella auripila (Winnertz) (Diptera: Sciaridae) and Megaselia halterata (Wood) (Diptera: Phoridae). Different numbers of adults emerged from each of these composts. Highest numbers of L. auripila emerged from spawned and pasteurized compost whereas lowest numbers of L. auripila emerged from full-grown compost. the emergence from full-grown compost was delayed, which could be explained by the delayed development of the larvae in this type of compost. High numbers of M. halterata emerged from compost that was completely colonized by the mycelia of the edible white button mushroom Agaricus bisporus (Lange) Imbach. The immediate covering of the compost with a casing layer significantly lowered the numbers of emerging M. halterata flies. Compared with the emergence pattern from full-grown and immediately-cased full-grown compost, adult M. halterata showed a delayed pattern of emergence in spawned compost. Adult M. halterata did not emerge from pasteurized compost. The results of these experiments enabled us to improve the timing of the application of insect pathogenic nematodes in the control of the larvae of both insect pests.     

Studies on an overwintering population of the mushroom phorid Megaselia halterata (Diptera: Phoridae) parasitized by Howardula husseyi (Nematoda: Allantonematidae)

An overwintering population of the mushroom phorid fly Megaselia halterata parasitized by Howardula husseyi was studied in an attempt to explain the winter decline in incidence of parasitism that has been observed in flies from mushroom farms. Fly larvae from eggs hatching in November developed into pupae in December and flies emerged in May. No selective mortality of parasitized specimens of larvae, pupae, or flies was observed. Dead parasites were found in only 10% of parasitized flies. The incidence of parasitism in the emerging flies (50%) was five times that of their parental generation and although parasitism significantly delayed fly emergence the delay was only 2–3 days. There was no evidence of winter decline in parasitism; instead there was strong evidence that parasitism enhanced phorid survival through the winter.     

Phoretic dispersal ofBrennandania lambi (Kcrzal) (Acari: Tarsonemida: Pygmephoridae) by mushroom flies (Diptera: Sciaridae and Phoridae) in New South Wales,Australia

The phoridMegaselia halterata (Wood) was found to carry significantly moreBrennandania lambi (Kerzal) than the sciaridLycoriella mali (Fitch) in samples collected from six mushroom farms.B. lambi was attached to both the phorids and mainly between the procoxae and mesosternum (57&) and between metacoxae and the first abdominal sternite (36%).B. lambi detached fromM. halterata only in the presence ofAgaricus mycelium. Evidence for dispersal ofB. lambi byL. mali andM. halterata is presented and the role of phoresy in the dispersal ofB. lambi is discussed.     

No evidence for behavioural adaptations to nematode parasitism by the fly Drosophila putrida

Behavioural adaptations of hosts to their parasites form an important component of the evolutionary dynamics of host–parasite interactions. As mushroom‐feeding Drosophila can tolerate deadly mycotoxins, but their Howardula nematode parasites cannot, we asked how consuming the potent mycotoxin α‐amanitin has affected this host–parasite interaction. We used the fly D. putrida and its parasite H. aoronymphium, which is both highly virulent and at high prevalence in some populations, and investigated whether adult flies utilize food with toxin to prevent infection in the next generation or consume the toxin to reduce the virulence of an already established infection. First, we found that uninfected females did not prefer to eat or lay their eggs on toxic food, indicating that selection has not acted on the flies to alter their behaviour towards α‐amanitin to prevent their offspring from becoming infected by Howardula. However, we cannot rule out that flies use an alternate cue that is associated with toxin presence in the wild. Second, we found that infected females did not prefer to eat food with α‐amanitin and that consuming α‐amanitin did not cure or reduce the virulence of the parasite in adults that were already infected. In sum, our results indicate there are no direct effects of eating α‐amanitin on this host–parasite interaction, and we suggest that toxin tolerance is more likely maintained by selection due to competition for resources than as a mechanism to avoid parasite infection or to reduce the virulence of infection.     

Phoretic relationship between the myceliophagous mite Microdispus lambi (Acari: Microdispidae) and mushroom flies in Spanish crops

We studied the role played by the phorid Megaselia halterata (Wood) and the sciarid Lycoriella auripila (Winnertz) in the phoretic dispersion of the myceliophagous mite Microdispus lambi (Acari: Pygmephoridae). Twenty‐four crops were monitored during 18 months in commercial mushroom farms in Castilla‐La Mancha (Spain). Adults of both species were collected weekly and the mites they carried were counted and identified. Both phorids (19.6%) and sciarids (4.4%) carried the mite M. lambi. The calculated load of each was 3.4 M. lambi mites per phorid and 1.9 per sciarid. The same percentage of male and female phorid was used as vector, but the load was slightly higher for females (1.86 mites per female compared with1.48 mites per male). A mean of 7.2% of the phorids examined in winter were vectors of M. lambi, while in spring and autumn of the first year the average was more than 22%. The mean load did not vary significantly between seasons. Inside the mushroom farms, less than 10% of a small initial population of phorids carried mites (less than two mites per phorid). As the cycle progressed, more than 35% of a larger population of emerging flies did so (average 3.5 mites per phorid vector). At the end of the growth cycle, the flies may fly off to colonise nearby farms, favouring the propagation of M. lambi from infested to uninfested crops. Megaselia halterata is the principal vector of M. lambi in the mushroom farms of Castilla‐La Mancha due to their high numbers, the high percentage carrying mites and the number of M. lambi they carry.     

Bendiocarb and diflubenzuron as substitute insecticides for endosulfan in commercial mushroom growing

Diflubenzuron and bendiocarb treatments of compost and casing soil at a rate of 1 g a.i./m² each, resulted in comparable or even improved compost and casing soil colonisation over endosulfan treatments (at a rate of 1.5 g a.i./m²). Alternating diflubenzuron/bendiocarb treatments were preferable to the reverse treatments, which tended to diminish total yield of mushrooms significantly by 1.4–3.9 kg/m². This effect was due to the casing soil treatment with diflubenzuron and not due to the compost treatment with bendiocarb after spawning. The effectiveness of these substitutes for endosulfan was tested on five commercial mushroom farms, where a similar or improved control of Megaselia halterata (Diptera: Phoridae) was obtained over control (endosulfan) treatments. If present, Lycoriella auripila (Diptera: Lycoriidae) was very effectively controlled by the substitute insecticides compared with endosulfan. Relatively high numbers of M. halterata were occasionally observed in endosulfan-treated houses, indicating that a certain level of resistance towards this insecticide may already have developed. This assessment of tolerance in M. halterata emphasised the need for substitute insecticides with different modes of action, in addition to environmental reasons. The chemicals should be alternately applied within individual crops to avoid resistance development.     

Host specificity of two pollinating seed‐consuming fly species is not related to soil moisture of host plant in the high Himalayas

Studying the drivers of host specificity can contribute to our understanding of the origin and evolution of obligate pollination mutualisms. The preference–performance hypothesis predicts that host plant choice of female insects is related mainly to the performance of their offspring. Soil moisture is thought to be particularly important for the survival of larvae and pupae that inhabit soil. In the high Himalayas, Rheum nobile and R. alexandrae differ in their distribution in terms of soil moisture; that is, R. nobile typically occurs in scree with well‐drained soils, R. alexandrae in wetlands. The two plant species are pollinated by their respective mutualistic seed‐consuming flies, Bradysia sp1. and Bradysia sp2. We investigated whether soil moisture is important for regulating host specificity by comparing pupation and adult emergence of the two fly species using field and laboratory experiments. Laboratory experiments revealed soil moisture did have significant effects on larval and pupal performances in both fly species, but the two fly species had similar optimal soil moisture requirements for pupation and adult emergence. Moreover, a field reciprocal transfer experiment showed that there was no significant difference in adult emergence for both fly species between their native and non‐native habitats. Nevertheless, Bradysia sp1., associated with R. nobile, was more tolerant to drought stress, while Bradysia sp2., associated with R. alexandrae, was more tolerant to flooding stress. These results indicate that soil moisture is unlikely to play a determining role in regulating host specificity of the two fly species. However, their pupation and adult emergence in response to extremely wet or dry soils are habitat‐specific.     

Susceptibility of mushroom pests to the insect-parasitic nematodes Steinernema feltiae and Heterorhabditis heliothidis

The potential of two species of insect-parasitic rhabditid nematodes (Steinernema feltiae, Heterorhabditis heliothidis) for biological control of mushroom flies was studied in pot trials. Three Diptera that commonly infest mushroom crops were used; the larvae of Megaselia halterata (Phoridae), Heteropeza pygmaea (Cecidomyiidae) and Lycoriella auripila (Sciaridae) were all susceptible to parasitism by both nematode species. Fewer adult phorids and sciarids emerged when compost was nematode-treated and, for L. auripila, the effects of nematode applications at spawning, casing or on both occasions were compared. Casing treatments were more effective than spawning treatments; little extra benefit was gained from applying the nematodes twice. Populations of paedogenetic larvae of H. pygmaea built up rapidly in untreated compost, but were reduced when S. feltiae was applied, and were eradicated by H. heliothidis. Because they can penetrate insect cuticle, as well as natural body openings, Heterorhabditis spp. may be more suitable than Steinernema spp. for the control of mushroom fly larvae.     

Survival and colonization of nematodes in a composting process

Nematodes are omnipresent in composts and are active in virtually all stages of the composting process. Major shifts in species composition, life strategies, and feeding behavior occur during the composting process. Due to the heat peak, nematodes can be virtually absent, but several taxa appear immediately when the temperatures drop. These comprise both taxa present before the heat peak and new taxa. However, it is not known how nematodes populate the compost. In this study, we aimed to assess the survival and colonization capacity of nematodes in compost. Our results showed that composting processes inaccessible to insects or not in contact with soil did not significantly influence nematode succession during composting. However, differences between treatments were found for some specific taxa (i.e., for Acrostichus sp., Neodiplogasteridae sp., Nygolaimoides sp., and Rhabditidae sp. 1), illustrating the importance of insects for the dispersal of nematodes to compost. Experiments in the lab with the blue bottle fly as a possible carrier demonstrated actual transport of nematodes isolated from compost by the fly (i.e., Halicephalobus cfr. gingivalis, Diploscapter coronatus, Diplogasteritus sp., Acrostichus sp., and Mesorhabditis sp.). Juveniles and dauer stages of Aphelenchoides sp., Panagrolaimus sp., and rhabditids survived an experimentally induced temperature peak, while members of Tylenchidae did not. In conclusion, our results indicate that the rapidly changing nematode community in compost is the result of both differential survival and colonization capacities.     

The biology of the phonotactic parasitoid, Homotrixa sp. (Diptera: Tachinidae), and its impact on the survival of male Sciarasaga quadrata (Orthoptera: Tettigoniidae) in the field

Abstract.

• 1 Unlike most parasitoids, tachinid flies of the tribe Ormiini use sound to locate their hosts. Although thought to exert selection pressure on their host's calling behaviour, little is known about the biology of ormiines. Accordingly, this study reports the biology and impact of the ormiine Hornotrixa sp. upon calling males of the univoltine bushcricket Sciarasaga quadrata Rentz in south-western Australia.
• 2 Populations of adult S.quadrata were monitored in the field over two successive calling seasons. Females, which do not call, were not parasitized by Hornotrixa sp., but the risk of parasitism for males increased as the 3-month calling season progressed. Parasitism did not commence until c. 2 weeks into the calling season, but by the end of the season up to 87% of surviving males were parasitized.
• 3 Parasitized males lived for 14 days and were found singing until their penultimate evening before death in the field. Unparasitized males lived on average 69 days and a maximum of 119 days.
• 4 Multiparasitism of hosts was common, with up to sixteen fly larvae found within parasitized males. The number of fly larvae within hosts significantly increased at the end of the season. However, successful emergence of fly larvae from hosts, as well as pupal size, significantly decreased as more than one fly larva developed within the host.
• 5 Hornotrixa sp. has a long pupal duration of 30–31 days at 20°C. As a consequence, only one complete fly generation, which overwinters in the pupal stage, is likely within each host generation.
• 6 No evidence for differential (size-bias) mortality by Hornotrixa sp. on male S.quadrata was found. The size of parasitized and unparasitized males collected in the field was not significantly different.
• 7 It is concluded that Hornotrixa sp. is a significant mortality factor acting on the survival of adult male S.quadrata.

     

Parasitoid wasp virulence

In nature, larvae of the fruit fly Drosophila melanogaster are commonly infected by parasitoid wasps. Following infection, flies mount an immune response termed cellular encapsulation in which fly immune cells form a multilayered capsule that covers and kills the wasp egg. Parasitoids have thus evolved virulence factors to suppress cellular encapsulation. To uncover the molecular mechanisms underlying the antiwasp response, we and others have begun identifying and functionally characterizing these virulence factors. Our recent work on the Drosophila parasitoid Ganaspis sp.1 has demonstrated that a virulence factor encoding a SERCA-type calcium pump plays an important role in Ganaspis sp.1 virulence. This venom SERCA antagonizes fly immune cell calcium signaling and thereby prevents the activation of the encapsulation response. In this way, the study of wasp virulence factors has revealed a novel aspect of fly immunity, namely a role for calcium signaling in fly immune cell activation, which is conserved with human immunity, again illustrating the marked conservation between fly and mammalian immune responses. Our findings demonstrate that the cellular encapsulation response can serve as a model of immune cell function and can also provide valuable insight into basic cell biological processes.     

Records of bizarre Jurassic brachycerans in the Daohugou biota,China (Diptera,Brachycera, Archisargidae and Rhagionemestriidae)

Abstract: Four new Brachycera fossils were collected from the Daohugou biota, China. Among these, two impressions demonstrating peculiar wing venation can be designated as two new species of a new genus (Mostovskisargus portentosus gen. et sp. nov. and M. signatus sp. nov.) referred to a new subfamily Mostovskisarginae (subfam. nov.) within Archisargidae. The third impression belongs to a new species (Calosargus (Pterosargus) sinicus sp. nov.) referred to the subgenus Pterosargus Mostovski, 1997 of Calosargus Mostovski, 1997 within Archisarginae, Archisargidae. It is only the second member of the subgenus Pterosargus worldwide. The fourth impression is a nearly complete fly, Jurassinemestrinus orientalis gen. et sp. nov. referable to Rhagionemestriidae. It reveals new morphological data about rhagionemestriids and extends the geographical distribution of this group beyond Europe and Central Asia during the Mesozoic. A correlation for the archisargid composition of taxa in Daohugou and Karabastau formations is discussed. The age of the fly‐bearing strata is briefly reassessed and can be more accurately limited to the Callovian–Oxfordian based on both biostratigraphical correlation and radiometric dating.     

A New Species of Dolichocephala,with a Key to the Species from China (Insecta: Diptera,Empididae)

Dolichocephala guangdongensis sp. n., a new aquatic dance fly (Diptera, Empididae), is described from Guangdong province in China. The relationships with its sister species are discussed. A key to the species of the genus Dolichocephala from China is presented.     

Fly pollination and duodichogamy in Bridelia stipularis and Cleistanthus sumatranus (Phyllanthaceae)

The globally distributed family Phyllanthaceae comprises 1745 species, all of which produce unisexual flowers, and most of the species occur in the tropics. The few species in which the pollination system has been studied are pollinated by flies, wasps, or specialized moths, and at least two species are duodichogamous. Here we report on the flowering phenology and pollination of the Chinese species Bridelia stipularis and Cleistanthus sumatranus, which were studied on the tropical Hainan Island. Both species are duodichogamous and indicated to be fly‐pollinated. The calliphorid species Chrysomya megacephala, Isomyia isomyia, Pierretia sp., Hemipyrellia sp., and Achoetandrus tufifacies were the most frequent flower visitors to B. stipularis; some of these species together with syrphids (Meliscaeva cinctella, Dideopsis sp., and Eristalis sp.) were the most abundant visitors to Cl. sumatranus flowers. Fruit set in hand‐pollinated flowers was higher than in open‐pollinated flowers, but because of natural fruit abortion the difference was not significant, indicating that fly pollinators do not limit reproduction of these Phyllanthaceae species. This study provides an addition to the limited Asian literature on pollination by flies and the even more limited literature on duodichogamy. It could help to broaden our understanding for analysis of the evolution of duodichogamy in future research, but the specific ecological factors behind the evolution of this rare sexual system are still difficult to generalize.     

A New Species of Chelipoda Macquart,with a Key to the Species from China (Diptera: Empididae)

Chelipoda nigraristata sp. n., a new aquatic dance fly of the subfamily Hemerodromiinae (Diptera, Empididae) is described from China. Relationships with other described species are discussed. A key to the species of the genus Chelipoda from China is presented for the first time.