Common Dung Beetles in Pastures of South-eastern Australia 9780643050907, 0643050906

Table of contents :
Contents
Acknowledgements
Why were dung beetles brought to Australia?
The CSIRO Dung Beetle Project
What does a dung beetle look like?
What do dung beetles do?
Distribution of dung beetles in Australia
Recognition of dung beetles
The spreading of dung beetles
Species location maps
Introduced dung beetles
Onitis alexis
Onitis aygulus
Onitis pecuarius
Onitis caffer
Onitis viridulus
Euoniiicellus intermedius
Euoniticellus africanus
Euoniticellus fulvus
Euoniticellus pallipes
Liatongus militaris
Onthophagus taurus
Onthophagus binodis
Onthophagus gazella
Onthophagus nigriventris
Sisyphus spinipes
Geotrupes spiniger
Hister nomas
Other beetles commonly found in dung pads
Native dung beetles
Beetles other than dung beetles
Maps showing potential and current distributions of species
Map 1 Onitis alexis
Map 2 Onitis aygulus
Map 3 Onitis pecuarius
Map 4 Onitis caffer
Map 5 Onitis viridulus
Map 6 Euoniticellus intermedius
Map 7 Euoniticellus africanus
Map 8 Euoniticellus fulvus
Map 9 Euoniticellus pallipes
Map 10 Liatongus militaris
Map 11 Onthophagus taurus
Map 12 Onthophagus binodis
Map 13 Onthophagus gazella
Map 14 Onthophagus nigriventris
Map 15 Sisyphus spinipes
Map 16 Geotrupes spiniger
Map 17 Hister nomas
Map 18 Onthophagus australis
Map 19 Onthophagus granulatus
Map 20 Onthophagus chepara and Onthophagus posticus
Map 21 Onthophagus auritus and Onthophagus dandalu
Map 22 Onthophagus capella and Onthophagus victoriensis
Map 23 Onthophagus pexatus
Map 24 Onthophagus pentacanthus and Onthophagus mniszechi
About the author

Citation preview

Common dung beetles in pastures of south-eastern Australia

Marina Tyndale-Biscoe

4111111~ C S I RO AUSTRALIA

Division of Entomology

National Library of Australia Cataloguing-tn-Publication Entry Tyndale-Biscoe. Marina. Common Dung Beetles in Pastures of South-eastern Australia ISBN 0 643 050906 1. Dung beetles - Australia. Southeastern. 1. CSIRO. Division of Entomology. II. Title.

caption for front cover figure: The largest of the introduced dung beetle species. Geotrupes spiniger.

© CSIRO Australia 1990.

Contents Acknowledgements ....................................................................................................v Why were dung beetles brought to Australia? ............................................................. 1 The CSIRO Dung Beetle Project ................................................................................... 1 What does a dung beetle look like? ............................................................................. 1 What do dung beetles do? ............................................................................................. 2 Distribution of dung beetles in Australia .................................................................. 3 Recognition of dung beetles ........................................................................................ 3 The spreading of dung beetles ..................................................................................... 4 Species location maps ................................................................................................ 4 Introduced dung beetles. Onitis alexis ...................................................................................................6 Onitis aygulus ................................................................................................8 Oniiis pecuarius ........................................................................................... 10 Onitis caifer ................................................................................................. 12 Onitis viridulus ............................................................................................ 14 Euoniiicellus intermedius ........................................................................... 16 Euoniticellus aJricanus ............................................................................... 18 Euoniticellus Julvus .....................................................................................20 Euoniticellus pallipes ..................................................................................22 Liatongus militaris ......................................................................................24 Onthophagus taurus ....................................................................................26 Onthophagus binodis ..................................................................................28 Onthophagus gazeZZa ....................................................................................30 Onthophagus nigriventris ...........................................................................32 Sisyphus spinipes ........................................................................................34 Geotrupes spiniger .......................................................................................36 Hister nomas ................................................................................................38

Other beetles commonly found in dung pads ........................................................... 40 Native dung beetles ....................................................................................... 40 Beetles other than dung beetles ................................................................... 41 Maps showing potential and current distributions of species ................................ 47 Map 1 Onitis alexis ...................................................................................48 Map 2 Onitis aygulus ................................................................................49 Map 3 Onitis pecuarius .............................................................................50

Maps showing potential and current distributions of species (continued) Map 4 Onitis caffer ................................................................................... 51 Map 5 Onitis viridulus .............................................................................. 52 Map 6 Euoniticellus intennedius ............................................................ 53 Map 7 Euoniticellus aJricanus ................................................................ 54 Map 8 Euoniticellus Julvus ...................................................................... 55 Map 9 Euoniticellus pallipes ................................................................... 56 Map 10 Liatongus militaris ....................................................................... 57 Map 11 Onthophagus taurus ...................................................................... 58 Map 12 Onthophagus binodis .................................................................... 59 Map 13 Onthophagus gazella .. ....................................................................60 Map 14 Onthophagus nigriventris ............................................................. 61 Map 15 Sisyphus spinipes .......................................................................... 62 Map 16 Geotrupes spiniger ......................................................................... 63 Map 17 Hister nomas ................................................................................. 64 Map 18 Onthophagus australis .................................................................. 65 Map 19 Onthophagus gmnulatus ............................................................... 66 Map 20 Onthophagus chepam and Onthophagus posticus ........................ 67 Map 21 Onthophagus auritus and Onthophagus dandalu ......................... 68 Map 22 Onthophagus capella and Onthophagus victoriensis ................... 69 Map 23 Onthophagus pexatus .................................................................... 70 Map 24 Onthophagus pentacanthus and Onthophagus mniszechi.. ......... 71 Detachable map overlay .................................................................. inside back cover

Acknowledgements The Dairy Research Council and the Australian Meat and Livestock Research and Development Corporation have, together with CSIRO, funded the publication of this booklet. I am grateful to Dr R. Sutherst. for generating the potential distribution maps of the introduced dung beetle species by the use of CLIMEX; to Mr John Green, for taking the photographs; and to Mrs Sandy Smith and Mr Chris Hunt for doing all the artwork and generally preparing the manuscript for publication.

C S I RO AUST RALIA Dairy Research Council

Division of Entomology

Australian Meat and Livestock Research and Development Corporation

v

antenna eye

~~~ j;,--c: I'~ " , pi) 5 ~~~

p" \

foreleg (frontal) horn on head pronotal lobe or hom

- - + - - - - pronotum

-'----'---'-~-;-~-

wing cover (elytra)

hind leg----=;-

Figure 1. Dorsal view of stylised dung beetle

Why were dung beetles brought to Australia? When Europeans created open pastures and introduced cattle. sheep and horses into Australia. they produced a new environment which few native dung insects were able to use. In particular. the dung of cattle was quite different from the fibrous. pellet-like droppings of native marsupials. Most Australian native dung beetles are adapted to living in woodlands and have little impact on the soft. moist dung pads of introduced stock in pastures. Nevertheless. a few native species in south-eastern Australia can, at times, be found in the dung of cattle and horses. Their activity is sporadic. however. and they are unable to conSistently bury the dung produced by the introduced animals. As a result. pasture pollution with dung. especially in heavily stocked areas, has become an important environmental and management problem. Undisturbed dung pads can smother the grass beneath them and promote an unpalatable growth of rank grass around them. Major fly pests breed in cattle dung: among these are the blood-sucking buffalo fly infesting cattle in the tropics, and the bush fly, a pest of man and other animals over most of Australia.

The CSIRO Dung Beetle Project. In the early sixties Dr G. F. Bornemissza suggested that, to solve this problem, dung beetles from countries with large. native herbivores be introduced into Australia. Subsequently, between 1968 and 1982 the CSIRO Division of Entomology imported over 45 species from Africa. Europe and Asia. Each species was reared in the laboratory under quarantine conditions for one or more generations before being released into climatically suitable areas. The benefits of dung burial are numerous. It cleans up pastures and replaces nutrients in the soil. Increased water penetration into the soil through beetle tunnels results in greater water retention and hence less runoff, decreasing soil erosion. Other benefits include better root penetration and soil aeration. Rapid dung burial destroys the breeding sites of the fly pests and reduces the free living stages of gastro-intestinal paraSites of livestock.

What does a dung beetle look like? Adult dung beetles are insects with a hard outside cuticle. The body has a head, a pronotum and an abdomen (Figure 1). Three pairs of legs are attached to the ventral (lower) surface of the body and there are folded wings lying over the dorsal (upper) surface underneath the wing covers. When not in flight the wings cannot be seen since the wing covers completely cover them. To fly, the beetles have to lift up the wing covers, and unfold the wings, which are generally transparent or whitish.

1

What do dung beetles do? Adults and larvae of dung beetles feed only on dung and therefore are not a threat to the environment. This is in contrast to some other beetles found in dung pads, (not dung beetles) which feed on vegetable and other matter besides dung, while others are carnivorous. The activity of adult dung beetles can break up the dung pad, and most species bury the dung in the ground for feeding and breeding. Usually one pair (male and female) work together in digging a nest in the soil under the pad. They take the dung down and form it into ball- or sausage-shaped objects called brood masses (Figure 2). Each brood mass contains one to several eggs, each in a small chamber. The eggs hatch into larvae which feed on the dung within the brood mass. The larvae grow through three distinct stages called instars, and the growth rate generally increases with higher temperatures. At the end of the third instar the larvae cease feeding. and in some species can stop development and stay inactive for many months. This enables them to survive unfavourable seasonal conditions. Eventually they change into non-feeding pupae and then into adult beetles. The size of the adult beetle remains unchanged throughout life; they do not grow with age. The newly formed adult beetles dig their way out of the ground and fly to fresh dung pads in which to feed and breed. The generation length varies with each species, and there may be one to several generations produced each year. The adults of most introduced species feed and breed throughout the late spring, summer and autumn. and die in winter. The larvae overwinter underground. A few of the introduced species have adults which feed and breed during both autumn and spring, having spent the winter inactive in the soil. They generally die in summer, but their larvae survive underground and emerge to form the new generation the following autumn. Beetles tend to be most numerous when a new generation emerges, often in response to rainfall. As many as 3000 adults of the smaller species have been counted in some pads at these times. They feed on the dung flUids, and mutual interference prevents any burial activity. On those occasions the pads are shredded and deSiccated until only dry loose chaff remains; this quickly weathers away. When tunnelling beetles are in breeding condition and are not overcrowded, their digging activities result in excavated soil being pushed to the surface to form small heaps or soil casts underneath and at the edge of the pad. The dung is taken down the tunnels from the underSide of the pad, often leaving the crust intact. Such a pad may appear untouched although it is hollow or filled with soil, and also weathers away quickly. In contrast, the one ball rolling species currently established in NSW cuts off lumps of dung from the edge or top of the pad, moulds them into balls which are rolled away and fixed to the bases of the stems of surrounding vegetation. The weight of a dung ball can be several times that of the beetle. In this species the immatures develop above ground, and there is no dung burial.

2

Birds often prey on dung beetles, as do foxes. Occasionally all the dung pads in a region will be fragmented by the predators in their search for these tasty morsels. However, dung beetles have evolved an ability to crawl down into their tunnels very rapidly when the dung pad is disturbed, and this way many escape from being caught. Dung beetles often have mites on them. The mites are carnivorous, feeding on small larvae and eggs in the dung pad. They also reproduce in dung. Dung beetles act as "aeroplanes", carrying the mites from one dung pad to another.

-I

pupa

larva

adult breaks out of dung ball and digs its way to the soil surface

young adult

\ fresh dung pat I

egg

I

~ soil casts tunnel filled with soil

tunnel often lined with dung brood mass containing several eggs

brood ball containing one egg

-

'--ball is attached to base of surrounding vegetation

Figure 2 .

)

-

ball is rolled away

ball is cut out of dung pad

Life cycle of the dung beetle. 3

Distnoution of dung beetles in Australia Of the species released. some 17 are known to be established in NSW. 4 in Victoria and 4 in Tasmania. Their natural rate of spread is slow. since in 10 years most species are still within areas less than about 100 Ian radius around the release sites. Many areas suitable for exotic dung beetles have not yet been colonised. In order to speed up their natural rate of spread. beetles can be collected from where they are established. at times of abundance. and released into other parts of their potential ranges. The potential distribution of each species has been determined by matching climatic data from localities where the beetles occur in their region of origin with those having an equivalent climate in Australia.

Recognition of dung beetles The aim of this booklet is to help to identifY dung beetles found in dung pads in south eastern Australia. Photographs of each species of introduced beetle are presented. together with some information on their biology. The maps show the potential distribution in south-east Australia of each introduced species, as well as its current distribution. The latter will. of course. change as each species spreads. Distribution maps and photographs of some of the native dung beetles and other beetle species commonly found in pastures are also included. To recognise which species are present. beetles should be sought. and picked out of one to two day old dung pads (by hand) for comparison with the photographs. Generally the beetles will be quite clean and easy to handle. If necessary. they can be washed in water and killed by submersion in methylated spirits. When identifying beetles. a magnifYing glass can be useful for recognising some of the finer detail. Males and females within each species sometimes look similar. but it is usually the male which has the easiest distinguishing features. In many species. though not all. the males have long. thin forelegs. while the forelegs of the females are shorter and have bigger. broader claws (eg. Figure 3a male. Figure 3d female). To enhance the chance of obtaining a male for identification. several specimens of each species should be collected for examination in the hope that a male will be included in the sample.

The spreading of dung beetles Farmers aware of large populations of introduced beetles on their properties may encourage other farmers to collect beetles for release elsewhere. This will have no noticeable effect on their own dung beetle numbers. Collections can be made by placing beetle-infested pads into containers with ventilated lids (such as mosquito netting). with some loose soil or vegetation under

4

and between the pads to ensure an adequate air supply for the beetles. Care has to be taken not to allow the containers to become overheated in direct sunshine or in closed vehicles. Alternately, beetle infested pads can be placed in a bucket of cold water. Within minutes all live dung beetles will float to the surface, from where they should immediately be scooped up and placed in a container half filled with some moist soil or sand and covered with a ventilated lid. This way they can be transported to the release site. Larger beetles can be collected individually by sifting through both the dung pad and the soil immediately under the pad, using a small spade or a garden fork. To release the beetles, the containers simply have to be emptied on to fresh dung pads in a paddock permanently stocked with cattle or horses. This is to ensure a source of fresh dung during the establishment phase. Several hundred beetles of each species should be released at a time, all at the same place: separate species may be released in different places. It may take three to five years before they breed up into large enough numbers to be noticed. By helping each species to achieve Its potential distribution we will greatly advance the time when the benefits of dung beetles can be enjoyed by the whole community.

Species location maps. The maps refer to the distribution of individual species of introduced dung beetles. Marked on each are the locations where that species is currently established (current distribution), and locations where the species would become established if it were released there (potential distribution). To help identify localities on the maps, there is a map overlay on the inside of the back cover with major towns marked on it. This should be torn out of the booklet, and laid over each map, so that distribution pOints can be located more accurately. Distribution maps of the common native species are included in the booklet: these, of course, cannot be spread any further since they already occupy all their potential ranges.

5

INTRODUCED DUNG BEETLES Onitis alexis (Figure 3a-e, Map 1)

Length: 15-20mm.

Colour: Pronotum dark brown, sometimes with a coppel)' or green sheen, wing covers light brown.

Special distinguishing feature: In males the hind femur has a single sharp spur (Figure 3e) while in the female there is no spur.

Distribution: The species occurs throughout the whole of the warmer, drier regions of Africa south of the Sahara, as well as in southern Europe. It occurs in summer, even and winter rainfall regions.

Introductions to Australia: Releases were made between 1973 and 1984 in NSW, QLD, NT and WA, and the species has become established in all these states. It was spread by CSIRO from the release sites to other areas and hence it now has a wide distribution in NSW.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads mainly at dusk and dawn; a few beetles fly during the night.

Biology: The adult life span during summer is approximately three months. A breeding female can lay up to three eggs a day. In moist, sandy soil brood masses are buried up to 25cm deep. Each sausage-shaped brood mass contains one to four eggs, and there are several brood masses in a nest consisting of 150-200g of dung. Development time from egg to adult during summer is a little over two months during winter up to 10 months and it can take over a year if conditions for the species are unfavourable. Although in warm climates the species can have several generations per year, in southern NSW it only has one, or at most two. In warmer regions the species over-winters both as adults and laIvae, but in the cooler regions it does so only as larvae. Figure 3

6

(a) male, dorsal view (c) male, side view

(b) male, ventral view (d) female, front view (e) hind femur of male showing spur

a

b

c d

e

Actual size 15-20mm

7

Onitis aygulus (Figure 4a-e. Map 2)

Length: 20-25rnm. Colour: Pronotum dark brown. sometimes with a green sheen; wing covers light brown.

Special distinguishing feature: In males the hind femur has a double spur (Figure 4e), while in the female there is no spur.

Distribution: The species occurs in the cooler. drier parts of South Africa.

Introductions to Australia: Releases were made between 1977 and 1982 in NSW. SA and WA and the species is now established at some of the release sites in all three states.

Activity period of adults: From spring to autumn. ceasing temporarily during hot, dry summers.

Flight time: Beetles fly to fresh pads at dusk and dawn.

Biology: In moist. sandy soils this species buries its brood masses about 16cm deep. Each sausage-shaped brood mass contains up to eight eggs. with several brood masses per nest consisting of 500-700g of dung. Development time from egg to adult during summer is 2.5-3 months. during winter up to 10 months. and this is greatly extended during unfavourable periods. There is one. possibly two generations a year. The species probably over-winters in the larval stage.

Figure 4

8

(a) male. dorsal view (b) male. ventral view (c) male. side view (d) female. front view (e) hind femur of male showing double spur

a

b

c d

e

Actual size 20-25mm

9

Onitis pecuarius (Figure 5a-e. Map 3)

Length: 15-2Omm. Colour: Dark brown to black.

Special distinguishing feature: In males the hind femur has a double spur (Figure 5eL while in the female there is no spur.

Distribution: The species occurs in the summer rainfall regions in South Africa.

Introductions to Australia: Releases were made between 1976-1979 in NSW and QLD. and the species has become established at some release sites in both states.

Activity period of adults: From late spring to autumn.

Flight time: Beetles fly to fresh pads at dusk and dawn.

Biology: The adult life span during summer is probably several months. This species takes up to 4 days to commence dung burial after arrival at a fresh dung pad. A female can then produce 1-2 eggs per day. The sausage-shaped brood masses generally contain 1 egg each. and several brood masses are joined together in a nest. In cooler climates the species probably over-winters in the laNaI stage.

Figure 5

(a) male. dorsal view (b) male. ventral view male. side view (d) female. front view (e) hind femur of male showing double spur.

(c)

10

a

c

b

d

e

Actual size 15-20mm

11

Onitis caffer (Figure 6a-e, Map 4)

Length: 15-2Omm. Colour: Black.

Special distinguishing feature: In males the hind femur has a serrated ridge (Figure 6e), while in the female the femur has no serration.

Distribution: The species occurs throughout the cooler summer, even and winter rainfall regions of South Africa, and has been recorded from Mozambique.

Introductions to Australia: Releases were made between 1979-1984 in NSW, QLD and WA. To date the species is only known to be established around Moruya, NSW. and at some sites in WA.

Activity period of adults: Autumn and early winter, and in winter rainfall areas the species is active again in spring.

Flight time: Beetles fly to fresh pads at dusk and dawn.

Biology: Adult life span is about 3-4 months. and a female can lay up to 2 eggs per day. In mOist. sandy soils brood masses can be buried more than a metre deep. Each is sausage-shaped and contains one to four eggs. There are several brood masses per nest, weighing 500-1000g. Development time from egg to adult in the field takes 10-12 months. Some individuals can slow down their development and remain as larvae for two, or even three years. There is only one generation a year. Over-wintering is in the larval and adult stages, but they survive the summers only as larvae.

Figure 6

12

(a) male, dorsal view (b) male, ventral view (c) male, side view (d) female. front view (e) hind femur of male showing serrated edge

a

b

c d

e

Actual size 15-20mm

13

Onitis viridulus (Figure 7a-e. Map 5)

Length: 18-23mm. Colour: Dark brown.

Special distinguishing feature: In males the hind femur has a double spur (Figure 7e). while in the female there is no spur.

Distribution: The species occurs in the moist, warm parts of Africa south of the equator.

Introductions to Australia: Releases were made between 1976-1980 in NSW. QLD and NT. and the species has become established at most release sites.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads at dusk and dawn.

Biology: Nests are made at a very shallow depth in the soil. often less than 10cm deep. Each sausage-shaped brood mass contains one egg. with several brood masses per nest consisting of 100-300g of dung. Development time from egg to adult during summer is 1.5-2 months. though during cold and unfavourable conditions they can take much longer to develop. There are probably several generations produced each year in the field. and over-wintering prQbably takes place in the lalVal stage.

Figure 7

14

(a) male. dorsal view (b) male. ventral view (c) male. side view (d) female, front view (e) hind femur of male showing double spur

b

a

c

d

e

Actual size 18-23mm

15

Euoniticellus intennedius (Figure 8a-e, Map 6)

Length: 7 -9mm. Colour: Medium brown.

Special distinguishing features: The male has a curved, blunt horn on the head (Figure 8e), while the female ha! no horn (Figure 8d). The ventral surface of the hind femur of both sexes i! uniformly pale (Figure 8b). The pronotum has symmetrical dark markings.

Distribution: The species occurs in the warm, moist parts of Africa, from South Afric2 through to the Sahel and Arabia.

Introductions to Australia: Releases were made between 1971 and 1975 in all the mainland states, and the species has become established at most release sites except in Victoria. This species was collected and redistributed by CSIRO from the release sites to many other suitable areas in the late 1970's and is now widely distributed throughout the warmer parts of Australia. The potential Australian distribution includes southern NSW, VIC and TAS; however all releases south of latitude 33°30' S (about the latitude of Sydney) have failed to establish. Further research may clarify the reason for this.

Activity period of adults: From spring to autumn, and during warm spells even in the winter.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span during summer is one to two months, and during this period a female can lay about 80-130 eggs. Brood balls are buried 0-15cm deep; during dry spells they are placed immediately below the dung pad to take advantage of any moisture there. Each brood ball is separate and pear shaped, weighing 3-3.5g and contains a single egg. Development time from egg to adult during summer is 4-6 weeks, but during winter it takes much longer. The species has up to 6 generations a year. Over-wintering takes place both in the adult and larval stages in the warmer regions, but only in the lalVal stage in the cooler parts of its range. Figure 8

16

(a) male, dorsal view (b) male, ventral view (c) male, side view (d) female, front view (e) side view of head of male showing horn

a

b

c

d

e

8 Actual size 7-9mm

17

Euoniticellus africanus (Figure 9a-d. Map7)

Length: 8-13mm. Colour: Light to dark brown.

Special distinguishing features: The ventral surface of the hind femur in both sexes is pale. with a darker area in the middle (Figure 9b). The pronotum and wing covers are speckled with dark markings.

Distribution: The species occurs throughout Mozambique. Lesotho and South Africa. It occupies a slightly cooler temperature range than does Euoniticellus intermedius.

Introductions to Australia: Releases were made between 1971 and 1977 in all the states except the NT. and the species has been recovered at some release sites in NSW and VIC.

Activity period of adults: From late spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: The adult life span is 2-3 months. Each brood pall is separate. rounded and contains a single egg. Development time from egg to adult during summer is 1-2 months. The species has up to 4 generations a year in the warmer parts of its range. Over-wintering is probably in the larval stage.

Figure 9

18

(a) male. dorsal view (b) male. ventral view (c) male. side view (d) female. front view

Euoniticellus ftdvus (FigurelOa-d. Map 8)

Length: 8-12mm. Colour: Light to mid-brown.

Special distinguishing features: The pronotum has no speckling. The wing covers may have darker brown markings. but no speckling (Figure lOa) such as occurs in Euoniticellus ajricanus (Figure 9a).

Distribution: The species occurs all through central and southern Europe and in countries around the Mediterranean basin.

Introductions to Australia: Releases were made between 1978 and 1982 in NSW. VIC. TAS. SA and WA and the species has become established in all these states.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span during summer is two to three months. Brood balls are buried about 20 cm deep in moist. sandy soil. more shallowly in loam. They are separate. pear shaped. each containing a single egg. Development time from egg to adult during summer is 5-8 weeks. much longer during winter. They can produce up to three generations a year. and they over-winter as larvae.

Figure 10

20

(a) male. dorsal view (c) male. side view

(b) male. ventral view

(d) female. front view

a

b

c

d

Actual size 8-12mm

21

Euoniticellus pallipes (Figure lla-d, Map 9)

Length: 9-12mm. Colour: Light to mid-brown.

Special distinguishing features: The pronotum and the wing covers both have obvious dark speckling. The ventral surface of the hind femur in both sexes is pale with a dark area in the centre (Figure llb).

Distribution: The species occurs throughout southern Europe, through Asia Minor, Turkistan, Iran, Mghanistan and India.

Introductions to Australia: Releases were made between 1977 and 1982 in NSW, SA and WA, and the species has been recovered in NSW and WA.

Activity period of adults: From spring to autumn.

Flight time: Adults fly to fresh pads during the day.

Biology: Adult life span during summer is two to three months. Brood balls are separate, pear shaped, each containing a single egg. Development time from egg to adult during summer takes 5-7 weeks, much longer during winter. There are at least two generations per year, and overwintering probably takes place in the larval stages.

Figure 11

22

(a) male, dorsal view (b) male, ventral view (c) male, side view (d) female. front view

a

Liatongus militaris (Figure 12a-c, Map 10)

Length: 8-1Omm. Colour: brown -black-yellow.

Special distinguishing features: The lateral and posterior edges of the pronotum are distinctly yellowish and lighter in colour than is the centre (Figure 12a). The wing covers have broken black stripes running lengthwise (Figure 12a,c).

Distribution: The species occurs in eastern Africa, ranging from Ethiopia to South Africa.

Introductions to Australia: Releases were made between 1968 and 1979 in NSW, QLD, WA and NT, and the species has become established in all states except in WA.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: In moist, sandy soil brood balls are buried 0-15cm deep. Brood balls are single, oval shaped, and contain one egg each. Over-wintering takes place mainly in the larval stage.

Figure 12

24

(a) female, dorsal view showing dark stripes on wing covers (b) female, ventral view (c) male, side view

a

b

c

.ctual size 8-lOmm

25

Onthophagus taurus (Figure 13a-f. Map 11)

Length: 8-1Omm. Colour: Black.

Special distinguishing features: Some (major) males have a pair of curved frontal horns extending backwards half way over the pronotum (Figure 13a.c.e). Other (minor) males have tiny horns projecting up from the head (Figure 13d). The females have no horns (Figure 13t).

Distribution: The species occurs in central and southern Europe and Asia Minor. ranging from Spain through to Morocco. Turkey and Iran.

Introductions to Australia: Releases were made between 1975 and 1984 in NSW. VIC. SA. WA and TAS. and the species has become established in all these states.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span is 2-3 months. Brood balls are buried up to lOcm deep in moist, sandy soil. They are separate. pear shaped. each containing a single egg. Development time from egg to adult during summer is 6-8 weeks. but it is much longer during winter. There are at least two generations per year. In the southern range of its distribution the species over-winters in the larval stages.

Figure 13

26

(a) male . dorsal view (b) male. ventral view (c) male. side view showing large horns on head (d) male. side view showing smaller horns on the head (e) head of male showing shape of horns (t) female. front view

Actual size 8-10mm

b

a

c

e

d

f

27

Onthophagus binodis (Figure 14a-d, Map 12)

a

Length: 11-13mm. Colour: Matt black.

Special distinguishing feature: The beetles have a lobe on the pronotum which is generally larger in the male (Figure 14a,c) than it is in females (Figure 14d).

Distribution: The species occurs in South Africa. where it is mostly restricted to areas with al annual rainfall of more than 500 mm.

Introductions to Australia: Releases were made between 1971 and 1981 in NSW, VIC. TAS. SA. WA and QLD and Norfolk Island. and the species has become established in all states excep in Queensland.

Activity period of adults: From late spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span during summer is about nine weeks. and a a female can lay 1-3 eggs per day. Brood balls are buried up to 20cm deep in moist sandy soil. They an separate. oval shaped. each weighing 6-7g. Development time from egg to adult during summer is between 4-6 weeks. during winter development is considerably slower. The species has several generations a year. In the south east over· wintering takes place in the larval stage. Onthophagus binodis has a preference for seoury dung and does well in spray irrigated pastures.

Figure 14

28

(a) male. dorsal view showing large lobe on pronotum (b) male. ventral view (c) male. side view showing smaller lobe on pronotum (d) female, front view

a

b

c

d

Actual size 11-13mm

29

Onthophagus gazeUa (Figure 15a-d. Map 13)

Length: 1O-13mm. Colour: Light to dark brown.

Special distinguishing features: Some males (major) have a pair of short. straight horns on the head (Figure 15a.c). the female has no horns (Figure 15d).

Distribution: The species occurs over much of the hotter. drier parts of Mrica south of the Sahara.

Introductions to Australia: Releases were made between 1968 and 1978 throughout Australia and Norfolk Island, and the species has become established in Norfolk Island and in all states except VIC. SA and TAS. It was collected and redistributed extensively by CSIRO in the early 1980s. and now has a wide distribution in the warmer parts of the continent. The potential distribution in Australia for this species includes southern NSW. VIC and TAS. However. many inland releases south of latituds 34° S (south of Cowra) have failed to establish. Further research may clarify the reason for this.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads from dusk to dawn.

Biology: Adult life span during summer is about two months. and one female can produce 1-2 brood balls a day. In moist. sandy soil brood balls are buried 20-25cms deep. They are oval shaped. containing a single egg each. Development time from egg to adult during summer is 3-5 weeks. during winter it is considerably longer There are several generations a year. and the species overwinters both in the adult and in the larval stages.

Figure 15

(a) male, dorsal view (b) male, ventral view male. side view showing horns (d) female. front view (e) head of male showing horns

(c)

30

a

b

c

d

e

Actual size 1O-13mm

31

Onthophagus nigriventns (Figure 16a-d. Map 14)

Length: 1O-12mm. males including the horn 16mm. Colour: Wing covers brown. pronotum and ventral surface black.

Special distinguishing feature: Some (major) males have a single large. forwardly directed horn on the from edge of the pronotum with a well developed lobe above it (Figure 16a.c): other males and females just have a small ridge in place of the lobe (Figure 16d).

Distribution: The species occurs in the high rainfall highlands of East Africa.

Introduction to Australia: Releases were made between 1974 and 1982 in VIC. NSWand QLD. and the species has become established at some sites in NSW and QLD. It has also been introduced. and become established. on Norfolk Island.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span during summer is 2-4 months. and the females lay about one egga day. Brood balls are single and oval shaped. Development time from egg to adult during summer is 4-7 weeks. There are at least 2 generations a year. and in the southern parts of its range the species probably over-winters in the larval stage.

Figure 16

32

(a) male. dorsal view showing horn on pronotum (b) female. ventral view (c) male. side view showing lobe over the horn (d) female. side view

a

b

c

d

Actual size males to 16mm UncI. hom)

33

Sisyphus spinipes (Figure 17a-c. Map 15)

Length: 9-11mm. Colour: Dark brown.

Special distinguishing features: The legs are extremely long and thin. The surface of the beetle is covered with little spines (Figure 17b.c). causing the dung to stick to it. This results in the beetles being almost invisible in the dung pad.

Distribution: The species occurs throughout central and eastern Africa. from Abyssinia to South Africa.

Introductions to AustraHa: Releases were made between 1972 and 1978 in QLD. NSW and NT. and the specie~ has become established in QLD and NSW. It has been collected and redistributed by CSIRO from the release sites to other areas in Queensland where it is now widespread.

Activity period of adults: From spring to autumn.

Flight time: Beetles fly to fresh pads during the day.

Biology: Adult life span during summer is two to three months. Under moist conditions females lay about one egg a day. Brood balls are spherical. weighing about 3-5g, and contain one egg each. These are rolled away from the pad by the parent beetle. and are attached to the bases of stems of surrounding vegetation. Development time from egg to adult during summer is 6-11 weeks. during winter over 6 months. and development takes place above ground. There are several generations a year, and over-wintering is in the larval stages.

Figure 17

34

(a) male or female. dorsal view (b) male or female. ventral view (c) male or female. side view

a

b

c

Actual size

9-11mm :~5

Geotrupes spiniger

a

(Figure lSa-d, Map 16)

Length: 20-25mm. Colour: Dorsal surface black, ventral surface blue.

Special distinguishing feature: The colour of the ventral surface of this beetle is a brilliant deep blue (Figure) lSb).

Distribution: The species occurs throughout Europe, the Balkans and through the Middle East as far as the Pakistani border.

Introduction to Australia: Releases were made in 1979 and 19S0 in NSW and TAS. and the species is known to be established in both states. though at only one site in Tasmania.

Activity period of adults: The autumn, early winter and again in the spring.

Flight time: Beetles fly to fresh pads at dusk and at dawn.

Biology: Adult life span is at least six months. During warmer days in autumn and spring a female can lay about 10 eggs per week. Nests are generally deeper than 20cm in moist. sandy soil. Brood masses are sausage-shaped, and contain one egg each. Several brood masses are joined together in a nest. Developmental time from egg to adult from spring to autumn is about 6 months: but eggs laid in the autumn take up to a year to develop in the field. (autumn to autumn). The species has only one generation per year. It over-winters both in the adult and larval stages. but probably oversummers only in the larval stage.

Figure IS

(a) male, dorsal view

(b) male. ventral view (c) male. side view (d) female, front view

36

b

a

c

d

37

Hister nomas (Figure 19a-c. Map 17)

Length: 7 -9mm. Colour: Shiny black.

Special distinguishing features: Body is very smooth. with little demarcation between the pronotum and the wing covers. Only a fine line identifies this junction (Figure 19a). The head is sometimes tucked under the pronotum and is then not visible from a dorsal (Figure 19b).

Distribution: The species occurs in southern Africa and was introduced to Hawaii.

Introduction to Australia: Releases were made between 1968 and 1975 in QLD. NSW. VIC. TAS and WA. but the species has been recovered from NSW and QLD only.

Activity period of adults: Spring to autumn in the cooler range of its distribution.

Flight time: Beetles fly to fresh pads during the day.

Biology: This is not a true dung beetle. although it lives in dung. Both the adults and the larvae are carnivorous. feeding on fly larvae and nematodes in the dung. Adults can live for up to 6 months. and lay eggs directly into the dung pad. They do not bury dung. During warm weather beetles can lay 1-2 eggs a day. and the larvae are cannibalistic. Development from egg to adult takes 4-6 weeks during summer. There probably are several generations per year.

Figure 19

38

(a) male or female. dorsal view (b) male or female. dorsal view with head retracted (c) male or female. side view

a

b

c

Actual size 7-9mm

39

OTHER BEETLES COMMONLY FOUND INi DUNG PADS a There is a whole array of insects and other organisms which colonise dun,g pads at different times of the year, including native dung beetles. These cannot be readily recognised as being native species (as distinct from introduced species without comprehensive keys to aid with the identification. Such keys are available in E.G. Matthews' "A Revision of the Scarabaeine Dung Beetles of Australia, I Trib Onthophagini" and "II Tribe Scarabaeini" Australian Journal of Zoolog Supplement No.9, 1972. and Supplement No. 24. 1974. respectively. Dung fauna also includes beetles other than dung beetles; very little is known about many of these. To aid with the recognition of some of these insects photographs of the most commonly found native dung beetles as well as other dung living beetles are shown here. together with distribution maps of each species where these are known.

NATIVE DUNG BEETLES Figure 20 (a) Onihophagus australis:

(b) (c) (d) (e)

Onthophagus australis: Onthophagus australis: Onihophagus granulatus: Onthophagus granulatus:

female and some (minor) males. black with green or coppery sheen. without horns (Map 18). some (major) males with horns. head of male showing horns. male. brown with black markings (Map 19). female.

Figure 21 (a) Onthophagus chepara: (b) Onthophagus posticus:

(c) Onihophagus auritus: (d) Onthophagus auritus: (e) Onthophagus dandalu: (f) Onthophagus dandalu:

40

sexes Similar. flattened. shiny black (Map 20). sexes similar. head and pronotum black. wing covers black with red stripes (Map 20). pronotum and wing covers shiny black. male has small horns (Map 21). head of male showing horns male. pronotum shiny green. with single lobe projecting forward (Map 21). female. pronotum shiny green. with two small bumps.

Figure 22 (a) Onthophagus capella:

(b) Onthophagus capella: (c) Onthophagus victoriensis:

(d) Onthophagus victoriensis: (e) Onthophagus pexatus:

female. black. two bumps on pronotum (Map 22). head of male with horns. black. anterior edge of head excavated (Map 22). Females without horns. horns on male short and bifurcated. sexes similar. small. matt black (Map 23).

Figure 23 (a) Onthophagus pentacanthus: (b) Onthophagus pentacanthus:

(c) Onthophagus mniszechi:

side view. one spine on head. four on pronotum. Sexes similar (Map 24). dorsal view. one spine on head. two on pronotum. Sexes similar (Map 24).

BEETLES OTHER THAN DUNG BEETLES Figure 24 (a) Aphodius tasmaniae: (b) Aphodius jimetarius:

(c) Aphodius pseudolividus:

(d) Proctophanes sculptus:

(e) Heteronychus arator:

elongated. dark brown to black. elongate. head and pronotum black. wing covers red. This species was originally an accidental introduction to Australia. elongate. light brown with dark brown shading in centre of pronotum and wing covers. This species was originally an accidental introduction to Australia. black. black. known as "Black beetle". This species was originally an accidental introduction to Australia.

41

a

b

c

Onthophagus australis

Actual size 11-13nun

d

e Onthophagus granulatus

Actual size 6-Smm

Figure 20 42

Current Distribution - the locations where the species were released and are established.

o 60

Potential Distribution - the locations where the species would become established if it were released there. The larger circles indicate more suitable conditions.

Current Distribution - the locations where the species were released and are established.

o

Potential Distribution - the locations where the species would become established if it were released there. The larger circles indicate more suitable conditions. 61

Current Distribution - the locations where the species were released and are established.

o 62

Potential Distribution - the locations where the species would become established if it were released there. The larger Circles indicate more suitable conditions.

Current Distribution - the locations where the species were released and are established.

o 64

Potential Distribution - the locations where the species would become established if it were released there. The larger circles indicate more suitable conditions.

Locatiolls wlIer(' tile llative species Ontlwphagus australis has becn recorded.

G5

Locatiolls wl]('I"(' the !latin' spccics Onl/lOp/W!}llS grnnlliCllw; has bcclI recorded.

(i(;

Locations where the native species Onthophagus chepara has been recorded. T

Locations where the native species Onthophagus posticus has been recorded.

67

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