Australian Dung Beetle Project
Encyclopedia
The Australian Dung Beetle Project (1965–1985), conceived and led by Dr. George Bornemissza
George Bornemissza
George Francis Bornemissza is a Hungarian-born entomologist and ecologist. He studied science at the University of Budapest before obtaining his PhD in zoology at the University of Innsbruck in Austria in 1950. At the end of that year he emigrated to Australia...

, of the Commonwealth Scientific and Industrial Research Organisation
Commonwealth Scientific and Industrial Research Organisation
The Commonwealth Scientific and Industrial Research Organisation is the national government body for scientific research in Australia...

 (CSIRO), was an international scientific research and biological control project with the primary goal to introduce foreign species of dung beetle to Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

 in order to control the polluting effects of cattle dung.

Background and Inception

Upon his arrival to Australia from Hungary
Hungary
Hungary , officially the Republic of Hungary , is a landlocked country in Central Europe. It is situated in the Carpathian Basin and is bordered by Slovakia to the north, Ukraine and Romania to the east, Serbia and Croatia to the south, Slovenia to the southwest and Austria to the west. The...

 in 1951, Dr. Bornemissza, an entomologist and ecologist, noted that Australian farmland was covered in a large number of cattle dung pads. This was in contrast to the fields of Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

 where the dung was removed and recycled back into the soil by various species of dung beetle
Dung beetle
Dung beetles are beetles that feed partly or exclusively on feces. All of these species belong to the superfamily Scarabaeoidea; most of them to the subfamilies Scarabaeinae and Aphodiinae of the family Scarabaeidae. This beetle can also be referred to as the scarab beetle. As most species of...

 (coprids). Native Australian species of beetle had co-evolved alongside marsupials such as the kangaroo
Kangaroo
A kangaroo is a marsupial from the family Macropodidae . In common use the term is used to describe the largest species from this family, especially those of the genus Macropus, Red Kangaroo, Antilopine Kangaroo, Eastern Grey Kangaroo and Western Grey Kangaroo. Kangaroos are endemic to the country...

 and wombat
Wombat
Wombats are Australian marsupials; they are short-legged, muscular quadrupeds, approximately in length with a short, stubby tail. They are adaptable in their habitat tolerances, and are found in forested, mountainous, and heathland areas of south-eastern Australia, including Tasmania, as well as...

, which produce small, hard, dry and fibrous pellets of dung. Cattle were relatively recently introduced to Australia by European settlers in the 1880s and produce large, soft, moist dung pads. Native beetles, with a few exceptions, are not adapted to utilise this type of dung as a food source or breeding ground and so without such fauna
Fauna
Fauna or faunæ is all of the animal life of any particular region or time. The corresponding term for plants is flora.Zoologists and paleontologists use fauna to refer to a typical collection of animals found in a specific time or place, e.g. the "Sonoran Desert fauna" or the "Burgess shale fauna"...

, the dung pads remain on the pasture
Pasture
Pasture is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep or swine. The vegetation of tended pasture, forage, consists mainly of grasses, with an interspersion of legumes and other forbs...

 and take months or even years to decompose. Cattle will not feed from the area of rank pasture surrounding the dung pad, and with the large quantity of dung produced (up to 12 pads per animal per day), this reduces the area of land available for cattle grazing by as much as 200 000 hectares per year. Cattle dung is also a primary breeding ground for several pestilent species of fly and parasitic worm. Bornemissza suggested in 1960 that the introduction to Australia of foreign dung beetle species, which had co-evolved alongside bovines and large herbivores, would be beneficial in removing the dung, thus improving cattle grazing and nutrient recycling and reducing the number of flies and worms.

Aims of the project

The overall aim of the Australian Dung Beetle Project was to establish a “minifauna”, that is, a subsection of the natural dung beetle fauna, of introduced dung beetles on the Australian mainland and in Tasmania
Tasmania
Tasmania is an Australian island and state. It is south of the continent, separated by Bass Strait. The state includes the island of Tasmania—the 26th largest island in the world—and the surrounding islands. The state has a population of 507,626 , of whom almost half reside in the greater Hobart...

. Once introduced, dung beetles in Australia were studied in order to determine their effects on:

1. Soil quality

Experiments were carried out in South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...

 to assess the rate at which dung beetles returned dung to the soil to aid nutrient recycling. It was found that dung beetles returned over 90% of the faecal nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

 excreted by each steer during the summer grazing season. Further experiments showed that the uptake of the essential minerals nitrogen, phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

 and sulphur in plants was over 80% greater in soils where dung beetles were active compared to those where they were not. In addition, tunnelling dung beetles assist root penetration and improve water infiltration to the soil, thereby reducing run-off
Surface runoff
Surface runoff is the water flow that occurs when soil is infiltrated to full capacity and excess water from rain, meltwater, or other sources flows over the land. This is a major component of the water cycle. Runoff that occurs on surfaces before reaching a channel is also called a nonpoint source...

 which can lead to the pollution
Pollution
Pollution is the introduction of contaminants into a natural environment that causes instability, disorder, harm or discomfort to the ecosystem i.e. physical systems or living organisms. Pollution can take the form of chemical substances or energy, such as noise, heat or light...

 of waterways

2. Fly control

Introducing dung beetles as a means to control fly populations was first utilised in Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...

 as a biological control against the horn fly (Haematobia irritans irritans). It was found that 96% fewer flies emerged from dung pads in open pastures with a dung beetle fauna in Hawaii compared to controls. It was surmised that the introduction of dung beetles to Australia would also have a reducing effect on the number of bush flies (bush fly (Musca vetustissima) and the blood-sucking buffalo fly (Haematobia irritans exigua). In addition to dung beetles, predacious beetles of the Histerid family, which are less sensitive to changes in soil moisture, were introduced to Australia as another means of controlling fly populations during periods of relative inactivity by dung beetles. However, these beetles were found to make little difference to the abundance of flies, so their introduction was discontinued in 1971.

3. Worm control

In experiments in Pretoria
Pretoria
Pretoria is a city located in the northern part of Gauteng Province, South Africa. It is one of the country's three capital cities, serving as the executive and de facto national capital; the others are Cape Town, the legislative capital, and Bloemfontein, the judicial capital.Pretoria is...

, dung beetles were shown to be effective at controlling infective worm populations breeding in dung. The activity of a complex of at least 20 species of coprid in cattle and sheep dung produced an average reduction of 85% of worms.

Care had to be taken to introduce only those species that would be most compatible with the Australian climates and soil types, that were not under great threat of predation or of themselves becoming pests, and that effectively dispersed dung pads within an ideal time frame of 48 hours so as to minimise successful fly and worm breeding by disrupting their reproductive cycles.

Quarantine

It was estimated that as many as 160 species of dung beetle would need to be imported into Australia in order to establish a minifauna of beetles that would be suitably adapted for the different Australian climates and soil types. However, it was also important that no other potential pests “piggy-backed” their way to Australia with the beetles. Therefore, strict quarantine
Quarantine
Quarantine is compulsory isolation, typically to contain the spread of something considered dangerous, often but not always disease. The word comes from the Italian quarantena, meaning forty-day period....

 measures were established to ensure no pestilent species made their way to Australia.

The first beetles to be imported to Australia came from Hawaii. Here, dung beetles, particularly the species Onthophagus gazella, had already been successfully introduced from Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 in order to help biologically control the numbers of horn fly. It was thought that these beetles would be a safe source since the only parasite present in Hawaii that does not already exist in Australia is the giant liver fluke, Fasciola gigantean, the eggs of which do not survive ingestion by beetles. Samples of O. gazella were shipped to the CSIRO research base in Canberra
Canberra
Canberra is the capital city of Australia. With a population of over 345,000, it is Australia's largest inland city and the eighth-largest city overall. The city is located at the northern end of the Australian Capital Territory , south-west of Sydney, and north-east of Melbourne...

, where they were kept under quarantine and observed before potential release. Unfortunately, direct importation from Hawaii was found to be unviable since it was discovered that the beetles were infested with potentially harmful pyemotid mites
MITES
MITES, or Minority Introduction to Engineering and Science, is a highly selective six-week summer program for rising high school seniors held at the Massachusetts Institute of Technology. Its purpose is to expose students from minority, or otherwise disadvantaged backgrounds, to the fields of...

. For this reason, these particular beetles were never released onto Australian pastures.

However, the mite-infested beetles were not destroyed and were instead used to breed new generations of beetles under sterile conditions. Dung beetle eggs were dipped in 3% formalin for 3 minutes to sterilise them, then packed into hand-rolled dung balls for incubation. These eggs successfully developed into adult beetles and were among those in the first batches released into the wild on 30 January 1968 in Lansdown, near Townsville, Queensland
Queensland
Queensland is a state of Australia, occupying the north-eastern section of the mainland continent. It is bordered by the Northern Territory, South Australia and New South Wales to the west, south-west and south respectively. To the east, Queensland is bordered by the Coral Sea and Pacific Ocean...

. This quarantine method was subsequently adopted at the CSIRO research base in Pretoria, South Africa, with the added step of the eggs being transported by air in sealed containers of sterilised peat moss.

First introductions

In the early years of the project, the dung beetle species O. gazella, which was able to remove dung pads in as little as 24 hours, showed the most promise in becoming established in Australia. After the first stocks were released onto pastureland in Northern Queensland in 1968, beetle recapture figures showed that the species had spread at the rate of 50–80 km per season, including the colonisation of two islands, Magnetic Island
Magnetic Island
Magnetic Island is an island offshore from the city of Townsville, Queensland, Australia. This mountainous island in Cleveland Bay has effectively become a suburb of Townsville, with 2,107 permanent residents. The island is accessible from Townsville Breakwater to Nelly Bay Harbour by ferry...

 and Palm Island
Palm Island
Palm Island may refer to:* Palm Island, Queensland, Australia, the community on Great Palm Island* Great Palm Island, Queensland, Australia, the island* Palm Island, Grenadines...

, 10 km and 30 km offshore respectively. By April 1970, O. gazella was firmly established over an area of 400 km2.

This performance however, was not uniformly excellent in all areas of Australia at all times of the season or year and so showed that Australia would need a number of other dung beetle species to fill the gaps in O. gazella’s activity.

The Australian Dung Beetle Project Research Unit in Pretoria

In order to find a suitable minifauna of beetles for Australia, Bornemissza set up the Australian Dung Beetle Research Unit in Pretoria, South Africa to find species that would complement the work of O. gazella. South Africa was chosen as a location to study dung beetles for several reasons. Firstly, there are a large number of different species from which to choose from (some 800 species south of the Zambezi
Zambezi
The Zambezi is the fourth-longest river in Africa, and the largest flowing into the Indian Ocean from Africa. The area of its basin is , slightly less than half that of the Nile...

 and Kunene
Kunene
Kunene may refer to:*Kunene Region, in Namibia*Kunene River, in Angola and Namibia*Kenny Kunene, a South African Businessman*Mazisi Kunene, a South African poet and historian...

 rivers alone), and these species have co-evolved alongside large bovids (e.g. African buffalo
African Buffalo
The African buffalo, affalo, nyati, Mbogo or Cape buffalo is a large African bovine. It is not closely related to the slightly larger wild Asian water buffalo, but its ancestry remains unclear...

) for many thousands of years. There are also homologies between the sub-tropical climates of areas of the two countries. The political climate in other parts of Africa and the more advanced nature of scientific research in South Africa also made it an ideal location. The Plant Protection Research Institute in South Africa funded much of the work at this research base. Further field stations were also maintained in Cape Town
Cape Town
Cape Town is the second-most populous city in South Africa, and the provincial capital and primate city of the Western Cape. As the seat of the National Parliament, it is also the legislative capital of the country. It forms part of the City of Cape Town metropolitan municipality...

 during 1978-1980 and in Hluhluwe Game Reserve during 1981-1986.

The aim was to find beetles that would match, as closely as possible, 8 selection criteria:

1. Genuine dung-breeders

Dung beetles for introduction to Australia needed to breed solely in dung and with no other food sources so as to avoid the species’ becoming pests in other habitats and to eliminate the possibility that they would avoid cattle dung altogether.

2. Predominantly bovine dung-breeders

Dung beetles tend to have a preference for a particular type of dung. To avoid the introduced dung beetles out-competing native Australian species of coprid that have a preference for other types of dung, the introduced species
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 needed to prefer cattle dung over other types.

3. Dung burial efficiency

The criterion was set that dung beetles selected for importation to Australia should be able to bury at least 25ml of dung per pair per day, on a pro-rata basis depending on beetle size and rate of activity.

4. Ease of handling

Beetles were chosen for export, which were able to breed easily in insectaries and could withstand strict the quarantine procedures imposed by Australian customs.

5. Fast breeder

Preference was given to dung beetle species that were able to reproduce quickly in order to maintain large stocks of beetles for release onto pastureland.

6. Compatibility

In order to minimise interspecific competition
Interspecific competition
Interspecific competition, in ecology, is a form of competition in which individuals of different species compete for the same resource in an ecosystem...

, dung beetles selected for export to Australia needed to be compatible with each other. Beetles that had co-evolved and co-existed (usually found on the same dung pads) were preferred.

7. Distribution range

Beetles adapted to a wide geographical range were preferred as this gave an indication as to how well they would establish across the range of climates found in the large Australian continent.

8. Taxonomy

Although not always possible, researchers preferred to select beetles that could be easily identified in terms of their taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 so as to be able to correctly identify them in mark, release and recapture follow-up experiments.

While most of this research was carried out in South Africa, it was recognised that further study in other areas of the world would be useful in selecting dung beetle species for introduction to Australian climates not matched by those in South Africa. To this end, a further research unit was set up in Montpellier
Montpellier
-Neighbourhoods:Since 2001, Montpellier has been divided into seven official neighbourhoods, themselves divided into sub-neighbourhoods. Each of them possesses a neighbourhood council....

, France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...

, as a base from which to study European species that may be more suited for introduction to cooler, southern areas of Australia.

Beetle collection surveys

Two types of field research
Field research
Field research is the collection of raw data in natural settings. It helps to reveal the habits and habitats of various organisms present in their natural surroundings...

 were carried out in Pretoria to identify beetles that would be suitable for export to Australia. Firstly, researchers took stocktaking trips to locations around South Africa in order to collect data on the biodiversity
Biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...

 and ecological abundance
Abundance
Abundance may refer to:In science and technology:* Abundance , the opposite of scarcities* Abundance , growing food with plentiful resources that will not run out -- sunshine, CO2, and waste or brine water....

 of the beetle fauna of an area, as well as the environmental conditions preferred by different species. Secondly, species earmarked as being potential candidates for biological control were then collected and taken back to the research base laboratory
Laboratory
A laboratory is a facility that provides controlled conditions in which scientific research, experiments, and measurement may be performed. The title of laboratory is also used for certain other facilities where the processes or equipment used are similar to those in scientific laboratories...

 where experiments were carried out to determine such things as beetle biology, habits, dung-disposing capacity, reproductive cycles and the ability to withstand quarantining procedures. It was necessary to devise and record methods of propagating dung beetles in large numbers, and this information forwarded onto the research base in Australia. In addition, extensive studies were carried out in the beetles’ native habitat to study their likely distribution patterns in Australia and thus aid in the selection of sites for their introduction. Later research has found that habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 specificity matching is one of the most important factors in determining whether or not an introduced species is successful in becoming established.

Samples of beetles that had already been successfully introduced to Australia were also collected for further study. This was important so as to enrich the gene pool
Gene pool
In population genetics, a gene pool is the complete set of unique alleles in a species or population.- Description :A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can survive bouts of intense selection...

 of beetle species within Australia and reduce the liability of any one species to become extinct because of a disease or chance mutation
Mutation
In molecular biology and genetics, mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or RNA sequence of a virus. They can be defined as sudden and spontaneous changes in the cell. Mutations are caused by radiation, viruses, transposons and mutagenic...

. Genetic variants were also imported to Hawaii in order to enrich the genetic diversity
Genetic diversity
Genetic diversity, the level of biodiversity, refers to the total number of genetic characteristics in the genetic makeup of a species. It is distinguished from genetic variability, which describes the tendency of genetic characteristics to vary....

 in that region, and as a thank you to the island for being cooperative in the research.

The CSIRO Dung Beetle Research base in Canberra

On arrival at the Canberra research unit, beetle eggs were transferred to dung balls and incubated. Adult beetles were then bred in insectaries for two or more generations in sterile conditions to eliminate the possibility of co-breeding parasitic mites or diseases endemic to Africa such as foot-and-mouth. Some beetle species were rejected at this stage and not subsequently released onto pastureland because difficulty was experienced in rearing those that entered a period of dormancy and some simply did not survive the stringent quarantining procedures.

Beetles that successfully survived and reproduced were despatched by air or road to cooperating farmers or CSIRO staff. They were packed into ventilated crates of damp peat and thousands at a time were simply tipped onto fresh dung pads at the chosen release sites. Most beetles were observed to instantly bury themselves into the dung. From then on, farmers helped to monitor beetle distribution and activity and carried identification cards in their vehicles so that they could monitor beetle activity.

The original review paper by Bornemissza states that by 1975, 23 species of dung beetle, including 3 genetic variants, had been released. By 1984, this number was 43 species, although it was found that 20 of these failed to establish. As well as the original “star” of the program, Onthophagus gazella, establishing well, other species such as Euoniticellus intermedius, Onthophagus binodis and Lissotes militaris were shown to have reproduced in their thousands and became abundant enough to allow the transfer of subpopulations of certain colonies to establish in new areas.

Further dung beetle research

The Australian Dung Beetle Project came to an end in 1986 when the Australian Meat Research Committee (AMRC), which then became the Australian Meat and Livestock Research and Development Corporation, withdrew their funding due to a shift in the focus of their work from on-farm production to off-farm marketing.

In 1998, interest in the project was revived when John Feehan, the manager of Soilcam, was invited by the Taroom Shire Landcare Group to give a seminar on the use of dung beetles. This led to Soilcam leading a two-week survey of the dung beetle fauna in south-eastern Queensland. The following December, groups of interested parties including graziers, scientists, government agencies and community groups met in Brisbane
Brisbane
Brisbane is the capital and most populous city in the Australian state of Queensland and the third most populous city in Australia. Brisbane's metropolitan area has a population of over 2 million, and the South East Queensland urban conurbation, centred around Brisbane, encompasses a population of...

 to discuss the future direction of dung beetle activities in Australia, and this in turn led to the formation of the National Dung Beetle Steering Committee, chaired by Mick Alexander. This committee proposed that a dung beetle project be undertaken in Queensland, which became a reality, thanks to funding from the National Heritage Trust, in October 2000.

The objective of the Queensland Dung Beetle Project was to provide the first comprehensive and quantitative
Quantitative
A quantitative property is one that exists in a range of magnitudes, and can therefore be measured with a number. Measurements of any particular quantitative property are expressed as a specific quantity, referred to as a unit, multiplied by a number. Examples of physical quantities are distance,...

 survey of the distribution and abundance of dung beetles across Queensland since the original CSIRO project, and to redistribute successful introduced species to other areas where they might be beneficial. This project was entitled “Improving Sustainable Land Management Systems in Queensland using Dung Beetles” and lasted for two years from January 2001 until December 2002.

The Queensland Dung Beetle project used pitfall traps at sites across Queensland to trap dung beetles, which were then identified and recorded. Out of the 29 species that were introduced to Queensland during the Australian Dung Beetle Project of 1965-1985, 15 of these were recaptured in 2001-2002. The most abundant and widely distributed of these were Onthophagus gazella and Euoniticellus intermedius. Some species, including Copris diversus and Onitis westermanni, which had been released during the original CSIRO project, were not recovered at all in the Queensland Dung Beetle Project, suggesting that they have not become established. Others, for example Onthophagus obliquus and Sisyphus infuscatus have been recovered since their original release but were also not found during this study. The project identified three species, namely Onitis caffer, Copris elphenor and Onitis vanderkelleni as candidates for redistribution to other areas of Queensland. These were chosen because, although they were not widely distributed across Queensland, they successfully established in the areas where they were found and had desirably rapid dung-burying capabilities.

The Queensland Dung Beetle Project concluded that one outcome of the study was to confirm the “outstanding success” of the original CSIRO project to select and introduce dung beetles into Australia, and “the impact of this on soil, water and pasture health, and on control of pest flies is undoubtedly worth many millions of dollars a year”. Further to this, the success of the Australian Dung Beetle Project is claimed to be the reason why Australians can now enjoy a café culture, as up until the 1950s, bush flies were so problematic that it was illegal for restaurants and cafés to offer outside dining unless a designated area was enclosed by fly-wire. A lesser effect has been had, however, on reducing the populations of Buffalo fly. This species of blood sucking fly remains a pest to livestock in Australia although they are present, but not pestilent in South Africa. Suggestions have been made that Australia needs further introduced species of predatory beetle (e.g. histerids) in order to help reduce the numbers of these flies.

A report by Penny Edwards of Landcare Australia in 2007 revealed that 23 species of beetle that had been introduced to Australia during the CSIRO project were still established species, whereas 20 had failed to establish (see table 1). The report recommended that in order to complete the work started by the Australian Dung Beetle Project, further action is required to:
  • identify geographical gaps in dung beetle distribution across the whole of Australia
  • identify seasonal gaps in dung beetle activity in all climate zones of Australia
  • clarify the role of native dung beetle species in the dispersal of cattle dung
  • undertake appropriate redistribution of introduced species
  • identify a suitable repository of unpublished dung beetle data
  • address the potential need for further introductions of dung beetles to Australia.


Table 1: Species of introduced dung beetle that have established in Australia (modified from a table in Edwards (2007) with some additional information from and ).
Key: ACT = Australian Capital Territory, NSW = New South Wales, NT = Northern Territories, QLD = Queensland, SA = South Australia, Tas = Tasmania, Vic = Victoria, WA = Western Australia)
Species Country of origin Total released First release Last release Areas of release Areas established Pasture type
Bubas bison France, Spain 1,613 April 1983 1996 (not by CSIRO) WA NSW, SA, WA Winter rainfall
Copris elphenor Klug South Africa 2,287 January 1977 May 1983 QLD QLD Summer rainfall
Copris hispanus Linnaeus Spain 294 October 1983 June 1994 WA WA Winter rainfall
Euoniticellus africanus Harold South Africa 49,009 October 1971 February 1984 NSW, QLD, SA, Tas, Vic, WA NSW, QLD Summer rainfall
Euoniticellus fulvus Goeze France, Turkey 76,944 March 1978 February 1983 NSW, SA, Tas, Vic, WA NSW, SA, Tas, Vic, WA Winter rainfall
Euoniticellus intermedius Reiche South Africa 248,637 November 1971 February 1984 ACT, NSW, NT, QLD, SA, Vic, WA NSW, NT, QLD, SA, Vic, WA Summer rainfall
Euoniticellus pallipes Fabricius Iran, Turkey 46,642 March 1977 September 1982 NSW, SA, WA NSW, SA, Vic, WA Winter rainfall
Geotrupes spiniger Marsham France 12,082 April 1979 December 1983 ACT, NSW, Tas, Vic ACT, NSW, SA, Tas, Vic Winter rainfall
Liatongus militaris Castelanu South Africa (via Hawaii) 70,450 January 1968 November 1979 NSW, NT, QLD, WA NSW, NT, QLD Summer rainfall
Onitis alexis Fabricius South Africa 186,441 August 1972 February 1984 NSW, NT, QLD, WA NSW, NT, QLD, SA, Vic, WA Rainfall, summer rainfall, winter rainfall
Onitis aygalus Fabricius South Africa 18,682 January 1977 January 1982 NSW, SA, WA NSW, SA, Vic, WA Winter rainfall
Onitis caffer Boheman South Africa 8,738 October 1979 April 1984 WA QLD, NSW, SA Summer rainfall, winter rainfall
Onitis pecuarius Lansberge South Africa 11,395 November 1976 May 1979 NSW, QLD NSW, QLD Summer rainfall
Onitis vanderkelleni Lansberge Kenya, Rwanda, Zaire 10,852 October 1974 February 1982 NSW, QLD QLD Summer rainfall
Onitis viridulus Bohemann South Africa 8,008 September 1976 July 1980 NSW, NT, QLD NSW, NT, QLD, WA Summer rainfall
Onthophagus binodis Thunberg South Africa 173,018 October 1971 February 1982 NSW, QLD, SA, Tas, Vic, WA, Norfolk Islands NSW, QLD, SA, Tas, Vic, WA, Norfolk Islands Winter rainfall
Onthophagus gazella Fabricius South Africa 420,415 February 1968 February 1984 ACT, NSW, NT, QLD, SA, Tas, Vic, WA, Norfolk Islands ACT, NSW, NT, QLD, SA, WA, Norfolk Islands Summer rainfall
Onthophagus nigiventris d'Orbigny East Africa 29,960 May 1975 March 1983 NSW NSW Summer rainfall
Onthophagus obliquus Nigeria, Senegal, Zaire 9,300 January 1976 November 1977 QLD, NT QLD Summer rainfall
Onthophagus sagittarius Sri Lanka (via Hawaii) 9,075 January 1968 March 1977 NSW, NT, QLD, WA NSW, NT Summer rainfall
Onthophagus taurus Schreber Spain, Greece, Italy, Turkey 164,499 February 1975 January 1984 NSW, SA, Tas, Vic, WA NSW, SA, Tas, Vic, WA Winter rainfall
Sisyphus rubrus Paschalidis South Africa 85,933 March 1973 February 1980 NSW, QLD, WA NSW, QLD Summer rainfall
Sisyphus spinipes South Africa 36,125 March 1972 December 1978 NSW, NT, QLD, WA NSW, QLD Summer rainfall

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