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.     

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.     

Populations of the bush fly (Musca vetustissima Walker) in arid Australia

Populations of the bush fly Musca vetustissima were usually low in arid south-western Australia. Herbaceous plants that grew only after rain resulted in changes to cattle dung that increased its attractiveness to bush flies for oviposition in laboratory tests. Larvae reared in such dung consistently produced large flies of high fecundity, and larval survival was usually high. Similar responses to the dung were evident from examination of flies sampled in the field. Although greatly increased bush fly abundance was possible only after major improvement in cattle dung, it was not a general phenomenon as immature survival did not always increase. Even major population increases resulted from periods of high immature survival that were brief relative to the duration of favourable dung. Nematodes Heterotylenchus sp. appeared to be an important mortality factor of the immature stages at such times. Build-up of high bush fly populations occurred only in cattle-grazing areas, but base-level abundance was similar regardless of the presence of cattle. Almost all female flies were gravid when cattle dung was unfavourable for breeding, and in areas without cattle. In arid areas, acceptable oviposition sites probably are more limiting than are sources of protein for oögensis.     

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.     

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.     

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.      

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.     

INSECTARY STUDIES ON THE CONTROL OF DUNG BREEDING FLIES BY THE ACTIVITY OF THE DUNG BEETLE, ONTHOPHAGUS GAZELLA F. (COLEOPTERA: SCARABAEINAE)

i
The Afro-Asian dung beetle, Onthophagus gazella F., buries bovine dung as food for its larvae so rapidly that when beetle populations are of the order of 4 insects per 100 c.c. dung, entire cow pads are completely broken up and buried within 30 to 40 hours.
Insectary studies show that this rate of dung disposal caused 80 to 100 per cent. reduction in the numbers of the bushfly Musca vetustissima Walker emerging from the pads. Surviving maggots gave rise to small, stunted flies of low or nil reproductive capacity. Viable fly eggs or maggots were never found in brood balls, and it is presumed that they were destroyed in the course of the elaborate process by which the beetles convert lumps of dung into brood balls.
Speed of dung burial is the critical factor in fly control. If half of a cow pad is buried within the first 24 hours, few or no adult flies emerge. More rapid burial within this period resulting in the complete removal of the cow pad, produces complete fly control.     

The prevalence of Stephanofilaria sp. in buffalo fly, Haematobia irritans exigua, in Central Queensland

Abstract  Buffalo fly ( Haematobia irritans exigua ) infestations of cattle are associated with characteristic lesions, the initial cause of which has been attributed to a filarial nematode of the genus Stephanofilaria , for which the fly acts as a vector. Survey work in the 1980s estimated the prevalence of microfilaria in female buffalo fly in Queensland at 2.91%. Since then no information has been published and the current prevalence of microfilarial infection in buffalo fly is not known. Buffalo fly were collected from four geographically distinct sites in Central Queensland in mid-summer 2004 and were dissected to estimate Stephanofilaria sp. infection rates . Larval stages of the nematodes were recovered from female flies from all four sites and the percentage of female flies from which nematodes were recovered ranged from 29% to 57%. The average number of larvae recovered from infected female flies ranged from 1.25 to 1.75. Whereas no infected male flies were found from Sites 2–4, larvae were recovered from 43% of male flies collected at Site 1. This high prevalence of filarial infection in buffalo flies implies a correspondingly high level of transmission to cattle in Central Queensland.     

Efficacy of cyromazine to control immature stable flies (Diptera: Muscidae) developing in winter hay feeding sites

Hay mixed with manure and urine residues at sites where hay has been provided as supplemental winter feed for cattle provide an excellent substrate for the development of immature stable flies, Stomoxys calcitrans (L.). Such sites are primary sources of early summer stable flies in the central United States and no effective measures are currently available to control fly development in them. A single application of granular cyromazine in May provided 97% reduction in the number of adult stable flies emerging from hay feeding sites. Stable fly control did not decline during the 12 wk season. A small decline in control was observed relative to anthomyiid, sarcophagid, and syrphid flies developing in the hay feeding sites. However, none of those flies are considered to be pests and > or = 50% control of those flies was maintained for 65 d after application. Cyromazine offers a safe and affordable option for the control of immature stable flies developing in winter hay feeding sites. Controlling those flies should reduce the estimated $2 billion per year of lost production in U.S. cattle industries attributable to stable flies.     

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.     

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.

     

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.     

The effects of land use change on native dung beetle diversity and function in Australia's Wet Tropics

The impacts of land use change on biodiversity and ecosystem functions are variable, particularly in fragmented tropical rainforest systems with high diversity. Dung beetles (Scarabaeinae) are an ideal group to investigate the relationship between land use change, diversity and ecosystem function as they are easily surveyed, sensitive to habitat modification and perform many ecosystem functions. Although this relationship has been investigated for dung beetles in some tropical regions, there has been no study assessing how native dung beetles in Australia's tropical rainforests respond to deforestation, and what the corresponding consequences are for dung removal (a key ecosystem function fulfilled by dung beetles). In this study we investigated the relationship between dung beetle community attributes (determined through trapping) and function (using dung removal experiments that allowed different dung beetle functional groups to access the dung) in rainforest and cleared pasture in a tropical landscape in Australia's Wet Tropics. Species richness, abundance and biomass were higher in rainforest compared to adjacent pasture, and species composition between these land use types differed significantly. However, average body size and evenness in body size were higher in pasture than in rainforest. Dung removal was higher in rainforest than in pasture when both functional groups or tunnelers only could access the dung. Increased dung removal in the rainforest was explained by higher biodiversity and dominance of a small number of species with distinct body sizes, as dung removal was best predicted by the evenness in body size of the community. Our findings suggest that functional traits (including body size and dung relocation behaviour) present in a dung beetle community are key drivers of dung removal. Overall, our results show that deforestation has reduced native dung beetle diversity in Australian tropical landscapes, which negatively impacts on the capacity for dung removal by dung beetles in this region.     

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.      

Summer decline in populations of the yellow dung fly: diapause or quiescence?

Abstract. Yellow dung fly ( Scathophaga stercoraria (L.)) populations on cow pastures in Central Europe usually show a characteristic summer decline in fly numbers. This has been related to their sensitivity to hot temperatures, but where and in what state the flies spend the summer has remained unclear. Field enclosure experiments revealed no evidence for survival over summer in the pupal stage, as adults never emerged in early autumn from eggs laid in late spring. Laboratory and field evidence shows instead that adults acclimatize physiologically by suppressing reproduction in favour of accumulating lipid (but not glycogen) reserves. Apparently they spend the summer in cooler, forested areas close to the pastures. As reproduction is not entirely shut down, it is suggested that this represents quiescence rather than diapause. Presumably this increases the survival of the flies during the hottest time of the season, and appears to be a flexible life-history strategy, particularly for late-born spring generation individuals.     

Superposition optics and the time of flight in onitine dung beetles

Dung beetles fly to fresh dung, with vision essential for flight navigation. The daily period of flight varies among different species: some beetles fly only in sunlight, others only when ambient light levels change rapidly during dusk or dawn and others in the constant dark of night. Measurements of the optical properties of the lenses, eye geometry and photoreceptor dimensions were used in a computer ray-tracing model to determine the optical performance of the superposition eyes of nine species of onitine dung beetles. Eye sensitivity to light is determined mainly by body size, by the refractive-index parameters and size of the crystalline cones, and by the photoreceptor dimensions. Based on the optics of the ommatidial lenses and absorption of light in the retina, the most sensitive eyes, found in the crepuscular-nocturnal beetles, are 85 times or nearly two log units more sensitive than the eyes of the diurnal beetles. Three possible criteria are considered to determine the best position for the retina: maximum amount of light absorbed in the target rhabdom; maximum amount of light falling on the target rhabdom (best focus); and maximum resolution. The structure and physiological optics of the superposition compound eyes of an onitine dung beetle are matched to the range of light intensities at which it flies. Accepted: 4 February 1998     

Dung beetles in pasture landscapes of Central America: proliferation of synanthropogenic species and decline of forest specialists

This study examines the spread of synanthropogenic dung beetles (species favoured by human activities) in pasture landscapes in Central America, and evaluates the role of forest fragments and regenerating patches of native vegetation in maintaining beetle diversity. Pitfall trapping was carried out at nine locations in El Salvador and seven in Atlantic Nicaragua that included both pasture and remnant or regenerating native vegetation. More dung beetle species occurred in forest fragments than in pastures. Community composition differed considerably between forest fragments from El Salvador and Nicaragua with many species restricted to either the Caribbean or Pacific regions. In contrast, dung beetle community composition and structure were largely the same in the pastures of El Salvador and Nicaragua, regardless of region or original habitat-type, and were similar to published results from pastures in Mexico and elsewhere on the Isthmus. Very small patches of native shrubs and tree stands (<2.5 ha) maintained no forest specialists in Nicaragua, whereas, in El Salvador, some forest specialists occurred even in the smallest stands of trees (ca. 0.25 ha). The study indicates that the expansion of cattle pastures has caused a regional decline in dung beetle diversity. Forest fragments and small isolated patches of native trees and shrubs maintain some of the diversity of the original landscape but their conservation value for dung beetles will depend on the biogeographical history of the sites.     

Landscape diversity of pasture dung beetle communities in the central region of mainland Japan and implications for conservation management

We studied the diversity of dung beetle communities in Japanese pastures to identify the factors that maintain or enhance the diversity of dung beetles at a landscape scale. We surveyed dung beetles from 17 pastures located in the northeastern part of Tochigi Prefecture, which is in the center of mainland Japan. From 1999 to 2001, surveys were conducted during the 6-month grazing period (May to October) by using dung baited basket traps. We also collected information about the environmental conditions and pasture management practices. Twenty-five dung beetle species belonging to Geotrupinae, Scarabaeinae, and Aphodiinae (including 13 tunneler and 12 dweller species) were recorded. The abundance of dweller species decreased with increasing elevation, possibly because of the effect of rainfall, whereas the species richness of tunneler species was affected by cattle disturbance and soil condition. Beetle species richness significantly increased with the number of years that the pastures had been grazed. Ivermectin administration did not appear to have any adverse effect on dung beetle abundance, species richness, or species diversity. The dung beetle datasets of the current study (including specific tunneler and dweller beetle groups) supported the widely documented positive relationship between local abundance and species distribution ranges. The within pasture, within area, and between area hierarchical additive partitioning of regional total diversity indicated that landscape-scale management should be implemented to conserve the regional diversity of the dung beetle communities inhabiting Japanese pastures.