Temperature-dependent development and parasitism rates of four species of Pteromalidae (Hymenoptera) parasitoids of house fly (Musca domestica) pupae

Parasitoid development, parasitoid-induced host mortality and parasitoid progeny emergence were determined at five constant temperatures for Muscidifurax raptor Girault and Sanders, Muscidifurax zaraptor Kogan and Legner, Spalangia cameroni Perkins and Spalangia endius Walker using pupae of the house fly, Musca domestica L., as hosts. At temperatures of 20, 25, 30 and 35 degrees C the median development times (days from oviposition to adult emergence), respectively, were M. raptor (28.4, 20.7, 14.3, 14.5), M. zaraptor (30.6, 22.8, 14.1, 14.2), S. cameroni (55.6, 35.2, 21.8, 25.0) and S. endius (52.4, 31.5, 16.3, 14.6). All species failed to emerge at 15 degrees C. Using densities of five parasitoids and 100 hosts and a 24 h exposure period, Muscidifurax species oviposited at a greater rate over a wider range of temperatures than Spalangia species. At 15, 20, 25, 30 and 35 degrees C the mean number of pupae killed per parasitoid were, respectively, M. raptor (1.4, 7.4, 10.5, 13.7, 14.1), M. zaraptor (0.0, 3.3, 8.9, 14.4, 15.0), S.cameroni (0.0, 7.8, 11.0, 11.9, 7.4), S.endius (0.6, 4.0, 7.5, 12.0, 11.7), and means of the number of parasitoid progeny per parasitoid were, respectively, M.raptor (0.2, 5.2, 7.9, 11.8, 11.6), M.zaraptor (1.3, 4.4, 8.2, 13.0, 13.7), S.cameroni (0.0, 2.4, 4.7, 5.1, 1.0), S.endius (0.0, 0.9, 3.4, 7.5, 4.9). Development and ovipositional activity in S.cameroni was strongly inhibited at 35 degrees C. The model by Sharpe & DeMichele (1977) was used to describe temperature-dependent development and the number of parasitoid progeny produced per parasitoid at temperatures of 15-30 degrees C in all species.     

Parasitism of house fly (Musca domestica) pupae by four species of Pteromalidae (Hymenoptera): effects of host–parasitoid densities and host distribution

Parasitoid-induced mortality of house fly, Musca domestica L., pupae and parasitoid progeny emergence by four species of pteromalid parasitoids, Muscidifurax raptor Girault & Sanders, M.zaraptor Kogan & Legner, Spalangia cameroni Perkins and S.endius Walker, were determined for a 24 h exposure period using parasitoid: host ratios ranging from 1:2 to 1:50. When the number of parasitoids was held constant (n = 5) and the numbers of hosts varied, and when the number of hosts was held constant (n = 100) and the number of parasitoids varied, both the number of pupae killed per parasitoid and the number of parasitoid progeny per parasitoid increased with increasing parasitoid:host ratios to reach an upper limit asymptotically. Maximum values were, respectively: M.raptor (14.7, 11.1), M.zaraptor (12.3, 9.3), S.cameroni (16.9, 5.5), S.endius (14.8, 9.7) with no consistent effects attributed to parasitoid interference. For M.raptor and S.cameroni at parasitoid:host ratios of 1:10, the pupal mortality and progeny emergence were determined for a 24 h exposure period when hosts were distributed in poultry manure at four levels of aggregation ranging from clumped to uniform. Pupal mortality was least in clumped distributions, while parasitoid progeny emergence was not significantly different.     

Pupal parasitoids (Hymenoptera: Pteromalidae) of muscoid filth flies in Israel

Pupal parasitoids of muscoid flies were collected monthly on three farms in southern Israel in preparation for an IPM programme for the control of filth-breeding flies. 50% and 25.5% of viable puparia collected during 1985 and 1986/87, respectively, were parasitized. Three species of Spalangia Latreille and two species of Muscidifurax Girault & Saunders (Hymenoptera: Pteromalidae) accounted for 82.0% and 94.2% of parasitoids recovered in the two seasons. Variation in the seasonal abundance and distribution of the leading parasite species are discussed in connection with their conservation and possible use for augmentative releases within IPM projects.     

Early season dispersal of Muscidifurax zaraptor (Hymenoptera: Pteromalidae) utilizing freeze-killed housefly pupae as hosts

The pteromalid wasp, Muscidifurax zaraptor Kogan and Legner, was released at three locations at a dairy in May before housefly and stable fly breeding had begun. Freeze-killed housefly pupae were placed adjacent to the emerging parasites at biweekly intervals for a 6-week period. Hosts placed out weeks 0 and 2 were heavily parasitized. Decreased parasitism in hosts placed out at week 4 suggested that many of the M. zaraptor had dispersed or died. High parasitism of hosts placed in the field at week 6 was the result of second generation parasites emerging from pupae placed out at week 0. Parasitism of freeze-killed housefly pupae placed 6 m and in the four cardinal directions from the release points was similar but lower than for hosts placed adjacent to the emerging parasites. The study demonstrated that emerging M. zaraptor readily utilized nearby freeze-killed housefly pupae but the availability of these hosts did not deter the parasites from searching for additional hosts.     

Survey of parasitoids (Hymenoptera: Pteromalidae) of house fly (Diptera: Muscidae) pupae from broiler-breeder poultry houses in northwest Arkansas

Parasitoids were collected from broiler-breeder poultry houses in northwest Arkansas using sentinel bags containing house fly, Musca domestica L., pupae. At least five species of Pteromalidae were collected over a two-year sampling period: Pachycrepoideus vindemiae (Rondani), Spalangia endius Walker, S. cameroni Perkins, Muscidifurax spp., and Nasonia vitripennis (Walker).     

Development of microsporidia-infected Muscidifurax raptor (Hymenoptera: Pteromalidae) at different temperatures

Muscididfurax raptor, a pupal parasitoid of house flies and other filth flies, is commonly infected with the microsporidium Nosema muscidifuracis. To determine the effects of infection on developmental time, uninfected and infected adult M. raptor were allowed to parasitize pupae of the house fly (Musca domestica) for 24 h. Exposed pupae of the two groups (infected and uninfected) were held at 15, 20, 25, 30, 32, and 34 °C with 75–80% relative humidity. Development of infected M. raptor was significantly longer at all temperatures than that of uninfected parasitoids, resulting in approximately 7% extensions of developmental times. Uninfected females completed development in 14.6, 19.6, and 30.4 days at 30, 25, and 20 °C, respectively, compared with 15.8, 20.7, and 32.3 days for infected females at these temperatures. The differences in developmental times provided narrow windows for isolating large proportions of uninfected M. raptor females for disease management programs. This window was greatest at 20 °C; 61% of the uninfected females emerged by day 30, at which time only 10% of the infected females had emerged.     

Characterization of microsatellite loci in the house fly,Musca domestica L. (Diptera: Muscidae)

The house fly, Musca domestica L., is a cosmopolitan species with a capacity to transmit human pathogens. Here, we report on the development of polymorphic microsatellite loci for house flies and present preliminary results from four house fly subpopulations from Manhattan, Kansas. Twenty‐four microsatellite loci were isolated and characterized using DNA enriched for repeat sequences. Forty individuals from four locations in Kansas were assayed to identify for polymorphic loci. Eight loci were polymorphic with the number of alleles ranging from three to six.     

Seasonal abundance of stable flies and filth fly pupal parasitoids (Hymenoptera: Pteromalidae) at Florida equine facilities

Beginning in November 2007 and continuing until December 2009, weekly stable fly, Stomoxys calcitrans (L.), surveillance was conducted at four equine facilities near Ocala, FL, by using alsynite sticky traps for adults and by searching immature developmental sites for pupae. Adult stable fly trap captures were highly variable throughout the year, ranging from 0 to 1,400 flies per trap per farm. The greatest adult stable fly activity was observed during the spring months of March and April, with weekly three-trap means of 121 and 136 flies per farm, respectively. The importance of cultural control measures was most apparent on the only farm with no reported insecticide use and the lowest stable fly trap captures, where an intense daily sanitation and composting program was conducted. A survey of on-site filth fly pupae revealed that 99.9% of all parasitoids recovered were Spalangia spp., consisting of Spalangia cameroni Perkins (56.5%), Spalangia nigroaenea Curtis (34.0%), Spalangia endius Walker (5.8%), and Spalangia nigra Latreille (3.7%). The implications of these findings are discussed.     

Efficacy and safety of catnip (Nepeta cataria) as a novel filth fly repellent*

Catnip (Nepeta cataria) is known for its pseudo‐narcotic effects on cats. Recently, it has been reported as an effective mosquito repellent against several Aedes and Culex species, both topically and spatially. Our laboratory bioassays showed that catnip essential oil (at a dosage of 20 mg) resulted in average repellency rates of 96% against stable flies, Stomoxys calcitrans (L.) and 79% against houseflies, Musca domestica (L.), respectively. This finding suggested that the application of repellent could be used as part of filth fly management. Further evaluations of catnip oil toxicity were conducted to provide a broad‐spectrum safety profile of catnip oil use as a potential biting and nuisance insect repellent in urban settings. Acute oral, dermal, inhalation, primary dermal and eye irritation toxicity tests were performed. The acute oral LD₅₀ of catnip oil was found to be 3160 mg/kg body weight (BW) and 2710 mg/kg BW in female and male rats, respectively. The acute dermal LD₅₀ was > 5000 mg/kg BW. The acute inhalation LD₅₀ was observed to be > 10 000 mg/m³. Primary skin irritation tested on New Zealand white rabbits showed that catnip oil is a moderate irritant. Catnip oil was classified as practically non‐irritating to the eye. In comparison with other U.S. Environmental Protection Agency‐approved mosquito repellents (DEET, picaridin and p‐menthane‐3,8‐diol), catnip oil can be considered as a relatively safe repellent, which may cause minor skin irritation.     

Host age and pathogen exposure level as factors in the susceptibility of the house fly,Musca domestica (Diptera: Muscidae) to Beauveria bassiana

Adult house flies, Musca domestica L., of four ages, <1, 3, 7, and 14 day post-eclosion, were exposed to three strains of Beauveria bassiana (P89, L90 and 447). Flies were exposed to moistened filter paper treated with either a low (1.57×10⁴ conidia/cm²) or high (1.57×10⁵ conidia/cm²) concentration of each fungal strain for 6 h. Strain 447 was superior to the two house fly-derived B. bassiana strains in inducing host infection and mortality. Significant spikes in infection and mortality occurred as early as 5 days post-exposure with higher concentration exposures acting more quickly. Few differences were observed in either infection or mortality among the four fly age classes. On Day 10 post-exposure, 77% of the high-concentration, 447-exposed flies were infected, compared with only 24% of the flies from the P89 low-concentration exposure. Potential applications of these results in integrated house fly management programs are discussed.     

Diploidy restoration in Wolbachia-infected Muscidifurax uniraptor (Hymenoptera: Pteromalidae)

Thelytokous reproduction, where females produce diploid female offspring without fertilization, can be found in many insects. In some Hymenoptera species, thelytoky is induced by Wolbachia, a group of cytoplasmically inherited bacteria. We compare and contrast early embryonic development in the thelytokous parthenogenetic species Muscidifurax uniraptor with the development of unfertilized eggs of the closely related arrhenotokous species, Muscidifurax raptorellus. In the Wolbachia-infected parasitic wasp M. uniraptor, meiosis and the first mitotic division occur normally. Diploidy restoration is achieved following the completion of the first mitosis. This pattern differs in the timing of diploidy restoration from previously described cases of Wolbachia-associated thelytoky. Results presented here suggest that different cytogenetic mechanisms of diploidy restoration may occur in different species with Wolbachia-induced thelytoky.     

Effects of host age, host density and parent age on reproduction of the filth fly parasite Urolepis rufipes (Hymenoptera: Pteromalidae)

Urolepis rufipes Ashmead, a pteromalid wasp, was recently discovered parasitizing house fly and stable fly pupae in eastern Nebraska dairies. Studies have been conducted on the biology of this parasite to evaluate its potential as a biological control agent of stable flies (Stomoxys calcitrans (L.] and house flies (Musca domestica L.). House fly pupae were suitable as hosts for U.rufipes at all ages; however, significantly higher parasitism occurred on host pupae aged 96-120 h. Parasite-induced mortality (host mortality without progeny production) was higher than for other pteromalid parasites of filth flies under similar conditions. Parasitism increased with parasite--host ratio at 20 degrees C; however, the opposite was noted at 30 degrees C for parasite--host ratios ranging from 5:50 to 50:50. Fly eclosion decreased as parasite--host ratio increased at 20 degrees C, and no host eclosion occurred at the highest parasite--host ratios (20:50 and 50:50) at 30 degrees C. Females produced an average of 18.6 female and 7.6 male progeny. 88% of the progeny were produced during the first 6 days post parental eclosion. The short life span, low progeny emergence rate and high per cent host eclosion, in comparison with other parasite species, suggests that the Nebraska strain of U.rufipes may not an effective biological control agent of house flies.     

Production of Filth Fly Parasitoids (Hymenoptera: Pteromalidae) on Fresh and on Freeze-killed and Stored House Fly Pupae

Laboratory experiments were performed to assess the effects of age, status (fresh versus freezekilled), and storage regime on the suitability of house fly, Musca domestica L. (Diptera: Muscidae) pupae as hosts for Muscidifurax raptor Girault & Saunders, M. raptorellus Kogan & Legner, M. zaraptor Kogan & Legner, Spalangia cameroni Perkins, Trichomalopsis sarcophagae (Gahan) and Urolepis rufipes (Ashmead) (Hymenoptera: Pteromalidae). Production of all species was maximized on pupae aged 24 + h post-pupation. Fresh pupae could not be refrigerated at 10°C or less, or at 15°C without a significant decline in their suitability as hosts. Although production of S. cameroni was essentially limited to the use of fresh house fly pupae, M. raptor , M. raptorellus , M. zaraptor , T. sarcophagae and U. rufipes could be reared on either fresh or freeze-killed pupae stored at - 20 °C for up to 6 months prior to parasitism. The suitability of freeze-killed pupae declined during storage when used for production of male and female M. raptorellus and M. zaraptor , and possibly for male T. sarcophagae . No other effects of storage on parasitoid production were detected. These results suggest that insectaries can stockpile fly pupae in freezers during times of overproduction for future use in mass-rearing M. raptor, M. raptorellus, M. zaraptor, T. sarcophagae and U. rufipes as biocontrol agents of filth flies.     

Larvicidal potential of the polyol sweeteners erythritol and xylitol in two filth fly species

The house fly, Musca domestica (L.) (Diptera: Muscidae), and the stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae), are two filth flies responsible for significant economic losses in animal production. Although some chemical control products target adults of both species, differences in mouthpart morphology and behavior necessitates distinct modalities for each. For these reasons, larvicides are an attractive means of chemical control. We assessed the potential of the polyol sweeteners erythritol and xylitol as larvicides to the house fly and stable fly. LC₅₀ values of erythritol against 2^nd instar larvae were 34.94 mg/g media (house fly) and 22.10 mg/g media (stable fly). For xylitol, LC₅₀ values were 74.91 mg/g media (house fly) and 41.58 mg/g media (stable fly). When given a choice, neither species showed a preference for ovipositing in media treated with either sweetener at various concentrations or in media without sweetener. Significantly lower development from egg to adult was observed when the 2^nd instar LC₅₀ equivalent of each sweetener was present in the media compared to controls. Erythritol and xylitol both have larvicidal qualities, however their effective concentrations would necessitate creative product formulation and deployment methods to control all stages of developing flies.     

Winter survival of nuisance fly parasitoids (Hymenoptera: Pteromalidae) in Canada and Denmark

Independent studies were performed in Canada and in Denmark to assess the survival of parasitic wasps (Hymenoptera: Pteromalidae) wintering in puparia of house fly, Musca domestica Linnaeus (Diptera: Muscidae). Data in Canada were collected for Muscidifurax raptorGirault & Saunders, M. raptorellus Kogan & Legner, M. zaraptor Kogan & Legner, Nasonia vitripennis(Walker), Spalangia cameroni Perkins, Trichomalopsis sarcophagae (Gahan) and Urolepis rufipes (Ashmead) in three microsites at an outdoor cattle facility in southern Alberta. Survival was highest for N. vitripennis, T. sarcophagae and U. rufipes, ranging from near zero to c. 7%. No survival was observed for S. cameroni. Daily mean values for ambient air temperature (DMAT) averaged about -3.5 degrees C during exposure periods. Data for Denmark were collected for M. raptor, S. cameroni and U. rufipes in a dairy barn and in a swine barn. Survival of M. raptor and U. rufipes was higher than that of S. cameroni in the dairy barn (DMAT = 8.6 degrees C), with the three species having similar survival in the swine barn (DMAT = 15.4 degrees C). In both studies, parasitoids in egg stages were least likely to survive. These results identify the potential for T. sarcophagae and U. rufipes to be commercialized for use in northern climates as biocontrol agents for nuisance flies, compare directly the cold-hardiness of commercialized species (i.e. all of the above species excluding T. sarcophagae and U. rufipes), and document the importance of microsite on winter survival.     

Sex ratios of Spalangia endius (Hymenoptera: Pteromalidae), in relation to current theory

Abstract. 1. Spulungiu endius Walker is a solitary parasitoid of house fly puparia.
2. The sex and size of S.endius was not related to host size.
3. In the laboratory the mean sex ratio of all offspring of nine groups, each comprising twenty females, was consistently female-biased (x = 83.5%, range 79–87%). The sex ratio in the field was less female-biased and showed greater fluctuation (61–75%). This may be a consequence of females laying male eggs before mating, some females remaining unmated, possible shorter adult life expectancy in the field than in the laboratory, and, perhaps, the presence of conspecific females.
4. The sex ratio of offspring of individual females varied from 66% to 100% females, and males were deposited early in the oviposition sequence.
5. Although a large number of fly puparia died before adult flies or parasitoids emerged (64.5%; n = 5874), there was no differential mortality of either sex.
6. Our results fit no general sex ratio hypothesis and we conclude that (i) the genetic nature of sex ratios in these insects needs careful examination, and (ii) the prevalence of female-biased sex ratios in solitary parasitoids needs investigation.     

Parasitism rates of Muscidifurax raptorellus and Nasonia vitripennis (Hymenoptera: Pteromalidae) after individual and paired releases in New York poultry facilities

Commercially reared parasitoids were released into three high-rise, caged-layer poultry houses; one house received only N. vitripennis Walker, the second house received only M. raptorellus Kogan & Legner, and the third house received an equal ratio of both species. Overall, house fly parasitism by M. raptorellus was never higher than 7% in any house. Most parasitism in the M. raptorellus release house was attributed to N. vitripennis. Parasitism of house fly pupae by M. raptorellus did not significantly increase during or after the 6-wk release period in the house that received both parasitoids. However, a depression in total parasitism was not detected when releases of the two species were made in this house.     

Prospects for improving biological control of olive fruit fly,Bactrocera oleae (Diptera: Tephritidae), with introduced parasitoids (Hymenoptera)

Olive fruit fly is a key pest of olive and consequently a serious threat to olive fruit and oil production throughout the Mediterranean region. With the establishment of Bactrocera oleae in California a decade ago, interest was renewed in classical (introduction) biological control of the pest. Here we discuss the prospects of identifying natural enemies of B. oleae in Africa and Asia that may help reduce B. oleae populations in California and elsewhere. Based on the current understanding of Bactrocera phylogenetics, early opinions that B. oleae originated in Africa or western Asia rather than the Mediterranean region or the Near East are taxonomically and ecologically supportable. Closely related to cultivated olive, the wild olive Olea europaea cuspidata is widely distributed in southern and eastern Africa, the Arabian Peninsula, and eastwards into Asia as far as southwestern China. Little is known regarding the biology and ecology of B. oleae in Africa and eastern Asia, especially in wild olives. While the diversity of parasitoids of B. oleae in the Mediterranean region is low and unspecialized, a diverse assemblage of parasitoids is known from B. oleae in Africa. Conversely, regions in Asia have remained largely unexplored for B. oleae and its natural enemies.     

Partial purification and characterization of phenobarbital-induced house fly cytochrome P-450

Two forms of phenobarbital-induced cytochrome P-450 were partially purified from the Rutgers diazinon-resistant strain of house fly using cholate solubilization, polyethylene glycol 6000 precipitation, and chromatography on DEAE cellulose. The preparation of highest purity had an absorbance maximum of 452 nm, a specific content of 10.0 nmol/mg protein, and an apparent molecular weight of 60,000 when examined by sodium dodecyl sulfate polyacrylamide electrophoresis. The yield of the highly purified cytochrome P-450 was 2–3%. This form contained proportionately less cytochrome P-420 than the original cholate solubilized microsomes, and is thus apparently more stable. A second form of cytochrome P-450 having a specific content of 0.50–0.89 nmol/mg protein was eluted from DEAE cellulose with a 0-0.25 M salt gradient. This is consistent with a previously reported elution pattern for Emulgen 913-solubilized house fly microsomes. Several methods of solubilizing house fly microsomes were examined. High salt, 2M KCI, in the absence of detergents effectively solubilized cytochrome P-450 (50–70% recovery) with little or no conversion to cytochrome P-(420).     

Microsatellite loci in the house fly,Musca domestica L. (Diptera: Muscidae)

Genomic libraries from house flies enriched for (CA)₁₅ and (CAG)₁₀ repeats were constructed by using biotinylated probes. Twenty‐five loci were isolated and evaluated for polymorphisms in wild flies representing two geographically diverse populations. Fourteen of 19 dinucleotide loci, and one of six trinucleotide loci were polymorphic. One hundred and twenty‐seven alleles were detected, 39 of which were private. Average number of alleles per polymorphic locus was 8.4 ± 2.5 and average heterozygosity was 72 ± 4%. F_ST by the private allele method was 0.73. Three of 15 loci showed significant heterozygote deficiencies, attributed to null alleles. Five of 15 loci were amplified in the face fly, Musca autumnalis.