Breeding of the bush fly,Musca vetustissima Walker,in an over-wintering area during winter and spring

Field experiments in an area of south-western Australia where bush fly (Musca vetustissima) adults occur permanently, showed that it overwintered by continuous breeding; but only two to three generations occurred between June and September. Low survival of eggs and larvae in the food (cattle dung) from May to August was associated with rainfall rather than low temperatures. High egg-adult survival occurred in late August; lower egg and larval survival in September and early October was attributed partly to dung fauna. Egg and larval survival was high in late October until December but predation or parasitization caused low survival of puparia outside the dung. Experimental and wild adult flies were largest and therefore most fecund in early spring; smaller wild flies in late spring appeared to result from larval competition for food. Simultaneous high egg-adult survival, high fecundity and rapidly decreasing generation times in late winter and early spring provide a basis for explaining the major increase in adult bush fly abundance observed in some areas in mid-spring.     

SEASONAL ABUNDANCE OF MUSCA VETUSTISSIMA WALKER AND OTHER CATTLE DUNG FAUNA IN CENTRAL AUSTRALIA

Abstract
Bush flies, Musca vetustissima attracted to a human, and arthropod fauna attracted to fresh cattle dung in 24 hours, were sampled every 2 weeks for 2 years (1980–1982) near Alice Springs in central Australia. Substantial rain fell in both summers, but it was more prolonged in the second. The bush fly occurred and bred throughout the year, although its abundance was usually low. The only major increase in bush fly abundance occurred after the first summer's rainfall caused the growth of new herbage. This was followed by increased feeding on dung by adult flies and intensified breeding. Bush fly abundance was low in the second summer, despite evidence of a long period of continuous breeding. A variety of dung-feeding and predatory beetles and also mites was almost always present, although numbers were usually low. The introduced dung beetles Euoniticellus intermedius and Onthophagus gazella numerically dominated samples. Dung fauna abundance also increased after significant rainfall, particularly in the second summer. Dissections of the dung beetles showed that they bred at these times.     

The seasonal distribution and characteristics of bush fly Musca vetustissima Walker populations in south-western Australia

In the agricultural region of south-western Australia the bush fly Musca ventustissima Walker was found to occur permanently in the north-eastern half but it died out during winter in the south-west. The abundance of bush flies increased rapidly throughout the overwintering zone in early spring as a result of local breeding. Flies then dispersed south-westwards, completely repopulating the region during October. Immigrant flies were large and therefore highly fecund, and of advanced reproductive maturity. Local breeding then rapidly increased the population. Maximum abundance occurred in mid-spring in the north and east and in early summer in the south and west. Abundance was greatest in the north where large overwintering populations occurred, and in the far south where repopulation occurred early in the seasonal cycle of pasture growth so that the more favourable cattle dung produced flies that were more fecund than their immigrant parents. The abundance of bush flies declined throughout the region in summer, to negligible levels near the south-west coast but, paradoxically, higher levels occurred inland where there were fewer cattle. On average, flies were smaller and reproductively younger in summer. In areas where irrigated pastures produced more favourable dung in summer, the patterns of abundance and other characteristics of flies did not differ substantially from those in comparable non-irrigated areas. The abundance and other characteristics of a bush fly population 10 km from a cattle-grazing breeding site were similar to those at the site, indicating that constant dispersal is the basis of the bush fly's ubiquity.     

SEASONAL CHANGES IN FAVOURABILITY OF CATTLE DUNG IN CENTRAL QUEENSLAND FOR THREE SPECIES OF DUNG-BREEDING INSECTS

b
The favourability of cattle dung from a native pasture near Rockhampton, Queensland, as a food source was tested monthly in the laboratory for 2 yr using 3 dung-breeding insects: the buffalo fly, Haematobia irritans exigua De Meijere; the bush fly, Musca vetustissima Walker; and a dung beetle, Euoniticellus intermedius (Reiche). Dung produced by cattle grazing on this pasture during the summer wet season yielded larger flies of both species and more broods from the dung beetle than dung from the same pasture in winter. When reared in summer dung, the buffalo fly almost attained its maximum recorded size but the bush fly and dung beetle reached ca two-thirds maximum recorded size and brood production respectively. Bush flies failed to breed in dung collected for 4 consecutive months in winter each year but survival of buffalo flies showed no seasonal trends.
The early response of the buffalo fly to improving dung quality in late winter/early spring gives it an advantage enabling its populations to increase earlier than those of its competitors, including the dung beetle, E. intermedius.     

Seasonal changes in characteristics of cattle dung as a resource for an insect in southwestern Australia

The value of cattle dung as a food resource for the bush fly Musca vetustissima (Walker) in the winter rainfall agricultural region of southwestern Australia was assessed by bioassay in the laboratory. The size (headwidth) of adult females was measured from flies reared on different samples of dung. Variation in size correlated with seasonal patterns of pasture growth, larger flies being produced during the growing season from autumn to spring. Size declined with senescence of annual pastures in late spring and early summer, occurring later in southern areas where the growing season was longer. After pasture senescence, dung from shorter growing season areas usually produced larger flies, apparently a result of the inverse relationship between digestibility of feed and length of growing season. Dung from irrigated perennial pastures never produced flies as large as that from annual pastures but generally high values were sustained during summer. Grazing of cereal stubble and feeding of hay in annual pasture areas during summer usually caused some increase in fly size. A spontaneous resurgence in the size of flies often occurred several weeks after pasture senescence and was attributed to more thorough digestion as a result of reduced intake of less palatable dry pasture.     

Effects of adding exotic dung beetles to native fauna on bush fly breeding in the field

Egg to pupal survival of bush fly,Musca vetustissima Walker, under field conditions was examined during 1987/88 in an area of south-eastern Australia that had not been colonised by exotic dung beetles. In pads of cattle dung containing only the native fauna, fly survival ranged from 0.3% to 12.5%. The addition of 2 species of exotic dung beetles,Euoniticellus fulvus (Goeze) andOnthophagus taurus (Schreber) to field pads, in numbers similar to those observed at the collection site, reduced fly survival to between 0.3% and 4.4%. Fly survival in the presence of the native and exotic dung fauna was sufficiently low to keep fly breeding below their mean replacement level of 3% for most of the season. Widespread dispersal and establishment of exotic dung beetles in south-eastern Australia, alongside the native fauna, should lead to long-term reduction of the bush fly problem.      

Influence ofOnthophagus gazella on hornfly,Haematobia irritans density in irrigated pastures

Haematobia irritans (L.) breeding in flood irrigated pastures of the lower Colorado Desert of southeastern California continues to remain unacceptably high during warm seasons (>1,000 adult flies per bovine head) despite the presence of moderately abundant populations ofOnthophagus gazella F. This study suggests that densities of > 40–70 adult beetles per dung pad and giving pronounced dung shredding activity, caused fly mortality of 38–56 %. The continued high abundance of adult horn flies on cattle suggests that at > 50% mortality, the pasture environment still produces sufficient flies to saturate cattle, although emigration might be reduced. Additional species of scarabs may be necessary to increase fly mortality. However, the dung drying activity of existingO. gazella significantly could interfere with resident staphylinid beetle breeding, which was significantly lower in pastures whereO. gazella reached densities of 40 per dung pad. Scarab beetle activity might also impede the introduction of superior predatory species for biological control.      

Effects of pyriproxyfen and buprofezin on immature development and reproduction in the stable fly*

The stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae), is one of the most significant biting flies that affect cattle. The use of traditional insecticides for stable fly control has only a limited success owing to the insect's unique feeding behaviours and immature development sites. A laboratory study was conducted to evaluate the effects of two insect growth regulator (IGR) products, pyriproxyfen and buprofezin, on the development of the immature stages of the stable fly and the effects of pyriproxyfen on oviposition and egg hatch. Both pyriproxyfen and buprofezin had significant inhibitory effects on immature development. The LC₅₀s of pyriproxyfen and buprofezin were 0.002 and 18.92 p.p.m., respectively. Topical treatment of adult females with different doses of pyriproxyfen had significant negative effects on both female oviposition and egg hatching when 1‐ and 3‐day‐old females were treated, and the effects were dose dependent. A significant reduction in the mean number of eggs laid was observed only at the highest pyriproxyfen dose (8 µg/fly) and egg hatch was unaffected by pyriproxyfen treatment when 5‐day‐old females were treated. Results from the present study indicate that pyriproxyfen has the potential to be used as part of an integrated stable fly management programme.     

Chemostimuli implicated in selection of oviposition substrates by the stable fly Stomoxys calcitrans

Horse and cow dung were tested as substrates for oviposition by the stable fly Stomoxys calcitrans (L) (Diptera: Muscidae) in laboratory cages. Odour alone from either horse or cow dung was sufficient to attract flies for oviposition. This was confirmed in wind tunnel experiments, where both horse and cow dung were shown to attract gravid stable flies. However, when S. calcitrans was offered a choice between these two oviposition substrates, flies always chose horse dung over cow dung, both when allowed to contact the substrates and when relying on dung odour alone. Analyses of volatile compounds emanating from horse and cow dung by gas chromatography linked antennogram recordings from S. calcitrans antennae revealed no differences in the chemostimuli released from the two substrates. The predominant chemostimulant compounds in both substrates were carboxylic acids (butanoic acid), alcohols (oct-1-en-3-ol), aldehydes (decanal), ketones (octan-3-one), phenols (p-cresol), indoles (skatole), terpenes (beta-caryophyllene) and sulphides (dimethyl trisulphide). Higher levels (20-40 p.p.m.) of carbon dioxide were recorded over horse dung compared with cow dung, a factor that may contribute to the preference exhibited by S. calcitrans for this substrate for oviposition.     

Nepetalactones from essential oil of Nepeta cataria represent a stable fly feeding and oviposition repellent

The stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae), is one of the most serious pests to livestock. It feeds mainly on cattle and causes significant economic losses in the cattle industry. Standard stable fly control involving insecticides and sanitation is usually costly and often has limited effectiveness. As we continue to evaluate and develop safer fly control strategies, the present study reports on the effectiveness of catnip (Nepeta cataria L.) oil and its constituent compounds, nepetalactones, as stable fly repellents. The essential oil of catnip reduced the feeding of stable flies by >96% in an in vitro bioassay system, compared with other sesquiterpene-rich plant oils (e.g. amyris and sandalwood). Catnip oil demonstrated strong repellency against stable flies relative to other chemicals for repelling biting insects, including isolongifolenone, 2-methylpiperidinyl-3-cyclohexen-1-carboxamide and (1S,2'S)-2-methylpiperidinyl-3-cyclohexen-1-carboxamide. The repellency against stable flies of the most commonly used mosquito repellent, DEET, was relatively low. In field trials, two formulations of catnip oil provided >95% protection and were effective for up to 6 h when tested on cattle. Catnip oil also acted as a strong oviposition repellent and reduced gravid stable fly oviposition by 98%.     

Are Dung Beetles Driving Dung-Fly Abundance in Traditional Agricultural Areas in the Amazon?

We evaluated the effects of different land-use systems on the ability of dung beetles to control the population of detritus-feeding flies. We tested the hypotheses that intensification of land use will reduce dung beetles richness, abundance and biomass and, consequently, their dung burial ability, affecting the interaction between dung beetles and flies and reducing its effectiveness as a natural biological control. In the Brazilian Amazon we sampled dung beetles, fly larvae and adults; and recorded the rate of dung removal by dung beetles across a gradient of land-use intensity from primary forest, secondary forest, agroforestry, agriculture to pasture. Our results provide evidence that land-use intensification results in a reduction of the richness, abundance and biomass of dung beetles, and this in turn results in lower rates of dung removal in the most simplified systems. We found no significant differences in the abundance of fly larvae between the different systems of land use. However, the number of adult flies differed significantly between land-use systems, presenting higher abundance in those sites with greater intensity of use (pasture and agriculture) and a lower abundance of adult flies in forested systems (primary and secondary forests, and agroforestry). Information-theoretic model selection based on AICc revealed strong support for the influence of land-use systems, dung removal rates and dung beetle abundance, biomass and richness on adult dung-fly abundance. Our results also reveal that dung beetles are not solely responsible for fly control and that other factors linked to land use are influencing the populations of these detritus-feeding insects.     

Role of bacteria in the oviposition behaviour and larval development of stable flies

Stable flies, Stomoxys calcitrans (L.), are the most important pests of cattle in the United States. However, adequate management strategies for stable flies, especially for pastured cattle, are lacking. Microbial/symbiont-based approaches offer novel venues for management of insect pests and/or vector-borne human and animal pathogens. Unfortunately, the fundamental knowledge of stable fly-microbial associations and their effect on stable fly biology is lacking. In this study, stable flies laid greater numbers of eggs on a substrate with an active microbial community (> 95% of total eggs oviposited) than on a sterilized substrate. In addition, stable fly larvae could not develop in a sterilized natural or artificial substrate/medium. Bacteria were isolated and identified from a natural stable fly oviposition/developmental habitat and their individual effect on stable fly oviposition response and larval development was evaluated in laboratory bioassays. Of nine bacterial strains evaluated in the oviposition bioassays, Citrobacter freundii stimulated oviposition to the greatest extent. C. freundii also sustained stable fly development, but to a lesser degree than Serratia fanticola. Serratia marcescens and Aeromonas spp. neither stimulated oviposition nor supported stable fly development. These results demonstrate a stable fly bacterial symbiosis; stable fly larval development depends on a live microbial community in the natural habitat, and stable fly females are capable of selecting an oviposition site based on the microbially derived stimuli that indicate the suitability of the substrate for larval development. This study shows a promising starting point for exploiting stable fly-bacterial associations for development of novel approaches for stable fly management.     

Effects of avermectin residues in cattle dung on yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae) populations in grazed pastures

The effects of avermectin exposure on natural populations of the yellow dung fly, Scathophaga stercoraria Linnaeus, were investigated at the field scale on farms in south-west Scotland. Pastures forming the focus of the study were grazed with either untreated cattle or cattle receiving standard, manufacturer-recommended treatment regimes of an avermectin product. Flies were sampled between April and July in 2002 and 2003 using dung-baited pitfall traps. Abundance and wing asymmetry in S. stercoraria populations were examined in relation to a range of environmental and management variables (including avermectin exposure, pasture management intensity, weather and season). Data used for abundance analyses were collected in fields where treated cattle had been dosed with either doramectin or ivermectin, while the data for the asymmetry analyses were from a subset of fields where treated cattle had been dosed with doramectin only. While abundance of S. stercoraria varied significantly between years and with season, there was no difference in their abundance between fields grazed by avermectin-treated or untreated cattle. Asymmetry was significantly higher in fly populations in fields grazed by doramectin-treated cattle, suggesting that exposure to doramectin during development could have imposed some degree of environmental stress. While these results suggest that exposure to doramectin residues in dung on grazed pastures may have sublethal effects on the insects developing in that dung, there was no evident avermectin effect on the abundance of adult S. stercoraria occurring in the pastures.     

Developmental sites and relative abundance of immature stages of the stable fly (Diptera: Muscidae) in beef cattle feedlot pens in eastern Nebraska

A 3-yr study was done to determine where and at what relative frequency stable fly, Stomoxys calcitrans (L.), immatures develop in beef cattle feedlot pens and the relationship of stable fly immature versus adult sample densities. Pens within feedlots were divided into five areas (the feed apron, back fence, side fences, mound, and the general lot); from each area, three core samples were randomly taken weekly. In 1986 and 1987, the feed apron yielded the most immature stable flies (62.5%). The mound and side fences yielded a significant percentage of flies (24.6 and 8.4%, respectively). There was a strong correlation (overall r = 0.86) between numbers of immatures and numbers of adults 2 wk later. In 1988, a drought year, low numbers of immatures were collected, and only one correlation between numbers of immatures and adults was significant. In all 3 yr, sample densities of stable fly immatures peaked and began to decline by midseason. Waste management along the feed apron and mound areas could significantly reduce stable fly populations. Sampling immatures from these areas could permit prediction of adult numbers.     

The Nematode Heterotylenchus autumnalis and Face Fly Musca autumnalis: A Field Study in Northern California

Helerolylenchus aulumnalis was found in six northern California counties surveyed, and the incitlence of nematode infection of face flies ranged from 4.7 to 43.8%. Intensive studies at a cattle ranch in Yuba County showed that population densities of the host and nematode infections were highest in flies from cow pats receiving full sun. Average host population density was 105.7 puparia per pat, and nematode infection averaged 38.6%. Pats in partial sun averaged 13.5 puparia and 13,1% nematode infection. No face fly was recovered from shaded pats. When data from pats first exposed during day or night were compared, no significant differences in host population density or nematode infection rates were apparent. Uninfected and superinfected flies were more frequent than predicted by a Poisson distribution.Infected and uninfected female flies of all ages captured on white sticky traps appeared to feed with similar frequency upon a creatny substance which was probably acquired from cattle, However, older infected females fed less on blood and more upon dung than older uninfected females. Percent nematode infection and host population densities were highest in spring and early summer, declined to a midsummer low, and then increased slightly. Both dung-reared flies and captured females showed similar trends in abundance anti infection rates. Regression analysis indicated that H. autumnalis may not be regulating face fly population density.     

Post-dispersal predation of Acacia farnesiana seeds by Stator vachelliae (Bruchidae) in Central America

Summary Post-dispersal seed predation by the bruchid beetle Stator vachelliae was investigated in Santa Rosa National Park, Costa Rica. This insect finds the seeds of the leguminous Acacia farnesiana in the feces of horses, deer, and ctenosaur lizards, the current major dispersers. Patterns of oviposition and pre-adult survival of beetles in the seeds were investigated in a series of experiments using fresh horse dung. S. vachelliae never minded into the dung balls, attacking only those seeds located on the surface. Fresh horse dung did not attract insects more readily than dry dung. The proportion of seeds attacked was not related to their density in a defecation, and was similar in three areas with different densities of the host plant. In a fourth area with no fruiting A. farnesiana shrubs all seeds survived insect predation. Bruchids attacked a greater proportion of seeds at 1 m than at 5 m from the edge of the shrub's crown. Seeds were mainly removed from horse dung by rodents with similar intensity in all areas and at both distances; this seed removal interfered with bruchid oviposition and probably with bruchid survival. S. vachelliae oviposited less frequently on seeds in dung fully exposed to sun. When oviposition on a dung pile was high, the distribution of eggs on the seeds was clumped, suggesting that some seeds were preferred to others. By the end of the dry season, bruchids stopped attacking the seeds. The results show that the fate of both seeds and bruchids is greatly influenced by the location and time of defecation.     

Asymmetric competition in cattle dung between two species of Onthophagus dung beetle and the bush fly, Musca vetustissima

Abstract.

• 1 Competition in cattle dung pads between two dung beetles, Onthophagus ferox Harold and Onthophagus binodis Thunberg, and the bush fly, Musca vetustissima Walker, was investigated in laboratory experiments, to determine why spring fly abundance in the field did not fall following the introduction of O. binodis.
• 2 At low beetle densities, the number of eggs laid by each species was reduced by the second species. A similar amount of dung was buried by each species alone or by both together.
• 3 At high beetle densities O. binodis egg production was substantially affected by each additional O.ferox, but O.ferox egg production was not affected by each additional O.binodis. Asymmetric competition occurred because O.ferox buried more dung than O.binodis, and a greater proportion in day 1 (pre-emptive dung burial).
• 4 O.ferox caused greater M. vetustissima egg-puparia mortality than O. binodis. Mortality mostly occurred in young M. vetustissima larvae less than 1 day old. Total egg-puparia fly mortality was correlated better with the dung buried on day 1 than dung buried on day 8 (pre-emptive dung burial). O.binodis did not add to fly mortality by O.ferox at high densities because of asymmetric competition between the beetles.

     

Determination of the toxicity of faeces of cattle treated with an ivermectin sustained-release bolus and preference trials using a dung fly, Neomyia cornicina

The toxicity of dung from cattle treated with an ivermectin sustained-release bolus was estimated in terms of ivermectin or ivermectin equivalents, using a laboratory bioassay with the dung fly Neomyia cornicina Fabricius (Diptera, Muscidae). The mortalities of flies measured 7 days after feeding for 24 h on dung containing known concentrations of ivermectin (between 0.125 and 1 g ivermectin per gram fresh dung) were compared with the mortalities of insects fed for 24 h on dung from cattle treated 21 days previously with an ivermectin sustained-release bolus. The toxicity of the bolus dung was equivalent to dung containing 0.66 g ivermectin per gram fresh dung. To determine whether insects could differentiate between control dung and dung from bolus-treated cattle, choice-chamber tests were carried out. There was no significant difference in the percentage of females that chose either dung type, suggesting that they were unable to distinguish the dung of bolus-treated cattle from control dung. Results are discussed in relation to the impact that bolus use can have on the insect fauna of cattle dung.     

Ecosystem services provided by dung beetles in Australia

Although there are nearly 500 species of native dung beetles in Australia, most are adapted to small, hard, dry, pelletised marsupial droppings and not to dealing with the large, moist deposits of cattle. In 1788, Governor Arthur Phillip arrived at Botany Bay with five cows, two bulls, 44 sheep and seven horses: this signalled major changes in Australia. Now there are about 27 million cattle, whose annual dung production has a dry matter content of about 42 million tonnes. Until CSIRO introduced exotic dung beetles in the 1960s, the dung of these herbivores sat on the soil surface, sometimes for years, locking up organic matter, smothering pasture and polluting waterways. CSIRO introduced 53 exotic dung beetle species, of which 43 were released to the Australian mainland between 1965 and 1985. Twenty-three of these have become established, many of which have reached the natural limits of their distribution. I consider the reason for the failure of the other 30 species to establish and briefly review previous contributions to examining the role of dung beetles in delivering ecosystem services, noting that much of the published literature concerns laboratory studies. New field data are then examined on the way in which introduced species are transforming dung communities and the ecosystem services they provide. The capacity of deep-tunnelling dung beetles to transform the soil profile is examined along with their effects on pasture production and the flow of nutrients from dung on pasture. The biocontrol capacity of dung beetle activity is considered in relation to the native bush fly, Musca vetustissima, the introduced buffalo fly, Haematobia irritans exigua, and dung-borne intestinal parasites (helminths and Cryptosporidium). The rationale for introducing additional species to Australia is considered.     

Effects of horn fly and house fly (Diptera: Muscidae) larvae on the development of parasitic nematodes in bovine dung

Laboratory studies were conducted to determine the effects of horn fly, Haematobia irritans (L.), and house fly, Musca domestica L., larvae on the development of a mixed population of parasitic nematodes in compressed and crumbled bovine dung. Fresh dung (100 g per sample) from a single calf passing trichostrongyle type eggs was infested with 150 horn fly or 150 house fly eggs. After 14-15 d, more horn flies and house flies had emerged from the compressed dung than from the crumbled dung, but more third stage parasitic nematode larvae were recovered from the crumbled dung containing either fly species than from dung containing no flies.