Bandicoots

(Peramelidae and Peroryctidae)

Class Mammalia

Order Peramelemorphia

Family Peramelidae and Peroryctidae

Thumbnail description
Small to medium-sized marsupials, with long, tapering snouts and short tails (most species); ears small to large, especially pronounced in rabbit-bandicoots; most species share a similar body form and are uniform in color, although some species have posterior barring or dorsal longitudinal stripes

Size
Head and body length ranges from 6.7–10.4 in (17–26.5 cm) in mouse bandicoot to 19.7–23.6 in (50–60 cm) in giant bandicoot; tail length from 4.3–4.7 in (11–12 cm) (mouse bandicoot) to 5.5–7.9 in (14–20 cm) (giant bandicoot); and weight from 4.9–6.5 oz (140–185 g) (mouse bandicoot) to 10.6 lb (4.8 kg) (giant bandicoot)

Number of genera, species
7 genera; 19 species

Habitat
Desert, grassland, woodland, forest, coastal complexes, rainforest, semi-urban

Conservation status
Peramelidae: Extinct: 3 species; Endangered: 1 species; Vulnerable: 3 species; Peroryctidae: Data Deficient: 7 species

Distribution
Peramelidae: Australia, New Guinea; Peroryctidae: New Guinea, Indonesia (Irian Jaya, Seram), Australia

Evolution and systematics

The order Peramelemorphia includes all the living bandicoots. They possess four or five pairs of blunt incisors in the upper jaw and three similar pairs in the lower jaw, and are thus polyprotodont. The hindfeet have the second and third toes joined in syndactyly. The order contains a single super-family, the Perameloidea, which is divided into two families: the Peramelidae contains all the non-spiny bandicoots and the pig-footed bandicoot (Chaeropus ecaudatus); the Peroryctidae includes the spiny bandicoots. While it appears that the order is intermediate between dasyuroids (polyprotodonts) and diprotodonts, the evolutionary origins of the bandicoots remain contentious and opinion varies, dependent on the significance given to dental or foot structure. Bandicoots may have evolved from dasyuroids, retaining polyprotodonty and separately evolving syndactyly, or from the diprotodonts, retaining their syndactyly and evolving polyprotodonty. But it is more plausible that they are derived from a proto-perameloid ancestor that produced two lines, one the terrestrial insectivorous/omnivorous bandicoots and the other the arboreal, herbivorous possums. Baverstock, et al. in 1990, suggested that this separation might have occurred around 48 million years ago (mya). Such fossil bandicoots as have been described differ little from modern forms.

The two families are discriminated by skull characteristics, that of peramelids being flattened in lateral view while the skull of peroryctids is more or less cylindrical. Other skull characters have been described by Groves and Flannery in 1990. The fur in many peroryctids is harsh and spiny. Peroryctids mostly inhabit rainforest, in contrast to the relatively dry habitats used by peramelids. It is possible that the now-extinct pig-footed bandicoot should be separated from the other species within the Peramelinae, as it exhibits a number of distinguishing characters of structure and behavior.

Physical characteristics

Bandicoots are small marsupials with a long, pointed snout, and are stockily built, with short limbs and neck. The ears are generally short and rounded, although more elongate in the genus Perameles. The pig-footed bandicoots have long, erect ears. The tail is thin and short in most species, although it is

long and crested in the pig-footed bandicoot. The teeth are small, relatively even in size, and sharply pointed. The dental formula is generally (I1-5/1-3 C1/1 P1-3/1-3, M1-4/1-4) but in Echymipera and Rhynchomeles there are only four pairs of upper incisors. The forefeet bear strong curved claws that are used in conjunction with the elongate muzzle to dig food items from the soil. Digits I and V are reduced in all species, and in Chaeropus only digits II and III are functional, giving rise to the common name. The hindfoot is elongate; digit I is reduced or absent, digits II and III are joined in syndactyly, digit IV is elongate, strong and powerful, while digit V is reduced or absent. Fur coloration is generally drab and unadorned, being darker on the dorsal surface and lighter ventrally. Exceptions exist, however. Perameles gunnii and P. bougainville have prominent pale posterior bars, Microperoryctes longicauda, M. papuensis, and Echymipera echinista are variously striped on the head and dorsum. The pig-footed bandicoot has a crested tail tipped with black.

Distribution

Bandicoots are confined to Australia, New Guinea, and the island of Seram. Prior to European settlement of Australia, bandicoots were widespread and at least one species was present in any given locality across almost the whole of the continent, in virtually all habitats. In the arid center of Australia, up to five species may have been found, whereas in tropical and temperate zones two or three species occurred. Some offshore islands are now the only places in which certain species survive. Similarly, bandicoots occur throughout New Guinea, from the coast to the central high mountain ranges, and some forms are restricted to off-shore islands. One species only occurs on Seram.

Habitat

All species are terrestrial. Australian habitats utilized by peramelids included: arid open-dense shrubland; sand plain, sand-ridge desert and spinifex grassland; temperate grasslands and grassy woodlands; wet and dry open-forest; deciduous vine thickets; heath and heathy woodlands and open-forest; savanna woodland, and shrubby grassland. Three species also use suburban gardens. In New Guinea, peroryctids occupy rainforest—lowland, primary and secondary highland; woodlands; subalpine grasslands; and are also found in gardens and regrowth forest. The Seram Island bandicoot was only found in tropical forest. Altitudinal range for bandicoots is from sea-level to 13,120 ft (4,000 m).

Behavior

Mutual avoidance is the predominant social behavior and most bandicoots are essentially solitary. During courtship and mating, male and female bandicoots associate for a limited time and several males may mate with a single female. The young may follow their mother for a short time after permanent emergence from the nest. Male-male interactions are always aggressive and in captivity will result in serious injury or death. Most species are nocturnal, some more strictly so than others, but southern brown bandicoots are often diurnal.

Bandicoots adopt several distinct postures when at rest, but when alert will often stand tripedally, with one foreleg raised and retracted towards the body, or stand erect on their hindfeet. Locomotion is quadrupedal and involves walking, running, galloping, and leaping. The latter is believed to be an escape mechanism. Vocalization is restricted to honks, snorts, and sneezes, which may be used to clear the nostrils after digging.

Feeding ecology and diet

Bandicoots are opportunistic and omnivorous, although the pig-footed bandicoot may have been more herbivorous. Most species obtain their food by first locating it through olfaction (and perhaps also by hearing) and then digging a conical pit to where the invertebrate or plant material is situated. The diet includes adult and larval insects (especially Coleoptera, Orthoptera, and Lepidoptera), earthworms, centipedes, seeds, bulbs, tubers, and hypogeous fungal sporocarps. Small vertebrates such as lizards and mice are occasionally eaten. In garden areas, and in tropical rainforest, fallen fruit is eaten.

Reproductive biology

The pouch opens to the rear and contains two crescentic rows of four nipples. Litters vary from one to five (average about two) in most species. The gestation period is very short—12.5 days in Perameles nasuta, P. gunnii, and Isoodon macrourus. These genera are polyestrous and the estrus cycle is about 20–25 days. Growth and development is rapid and in some species sexual maturity may be reached at theee to four months of age. Bandicoots may breed throughout the year, although some degree of seasonality is shown. Such seasonality may be dependent on climatic conditions; for example, eastern barred bandicoots in Tasmania do not breed during the coldest winter months, and the same species on the mainland ceases breeding during periods of drought. Breeding may be initiated by an increase in food availability (perhaps related to rainfall events), rates of change in temperature, or photoperiod. Mating is probably either polygynous or promiscuous.

One of the most significant features of bandicoot reproduction is the presence of a functional chorioallantoic placenta in addition to the yolk-sac. The placenta has evolved independently and is probably correlated with the rapid rate of development in bandicoots.

Bandicoots are short-lived—a maximum of two to three years in the wild, although they may reach five years of age in captivity.

Conservation status

Peramelidae: Of the 10 modern species, two are extinct. These are species from the arid interior—the pig-footed bandicoot (Chaeropus ecaudatus) and desert bandicoot (Perameles eremiana). The mainland form of the eastern barred bandicoot (P. gunnii) is Critically Endangered; the Tasmanian form is Vulnerable. One species, the western barred bandicoot (P. bougainville) is Endangered—it is extinct on the main-land and occurs only on Bernier and Dorre Islands in Shark Bay, Western Australia. Four subspecies are Vulnerable—the mainland and Barrow Island forms of the golden bandicoot, Nuyts southern brown bandicoot, and the Tasmanian form of the eastern barred bandicoot. Among the other forms (mainly subspecies) at least three are Near Threatened.

Peroryctidae: Little is known about the status of most species. One species, the Seram Island bandicoot (Rhynchomeles prattorum), is known only from the type series collected in 1920. Several other species, including the mouse bandicoot (Microperoryctes murina), David's echymipera (Echymipera davidi), Menzies' echymipera (E. echinista), and Papuan bandicoot (Microperoryctes papuensis), are rarely encountered and may be Vulnerable.

Significance to humans

Probably all species of peramelids were used as food by native Australians. The larger peroryctids are still hunted for food in New Guinea. Minor annoyance in suburban areas is caused by bandicoots digging foraging holes in lawns.

Species accounts

List of Species

Eastern barred bandicoot

Perameles gunnii

subfamily

Peramelinae

taxonomy

Perameles gunnii Gray, 1838, Tasmania, Australia.

other common names

English: Barred bandicoot, Tasmanian barred bandicoot, striped bandicoot, Gunn's bandicoot; German: Tasmanien-Langnasenbeutler.

physical characteristics

Head and body length is 10.6–13.8 in (270–350 mm); weight is 26.5–35.3 oz (750–1,000 g). Grayish brown fur with light stripes on its hindquarters. Large ears, tapered nose, and short tail.

distribution

Victoria, Tasmania, and formerly South Australia, Australia.

habitat

Grassland and grassy woodland, pasture, also gardens in suburban areas.

behavior

Nocturnal, solitary except when courting or mating or females with young.

feeding ecology and diet

Food is mainly obtained by digging after locating food items by smell. Small pits are dug using the forefeet and the long nose. Food is extracted and deftly manipulated in the front feet. Eats earthworms, adult and larval insects, other invertebrates, tubers, bulbs, and fallen fruit.

reproductive biology

Capable of breeding year-round but may cease in colder winter months at lower latitudes (Tasmania) or during hot, rainfall-deficient summers on the mainland. Gestation period 12.5 days, polyestrous, estrus cycle about 26 days. Chorioallantoic placenta formed at about 9.5 days of gestation and is retained in the uterus after parturition. Litter size one to five, average two to 2.5. Pouch life about 55 days, weaned at 70–80 days. The nest is a grass and leaf-lined scraped depression. Growth is rapid and sexual maturity may be reached at about four months. Sequential litters may be born throughout the female's two to three year lifespan. Mating is probably promiscuous.

conservation status

The mainland form is Critically Endangered. It only occurs in minuscule numbers at one site in the wild. A recovery program, involving reintroduction to protected sites of captivebred animals has been in operation since 1989. The principal continuing threat is predation by introduced carnivores, particularly red foxes and cats, for which species continuing control is essential for the reintroduced populations to survive. The Tasmanian population appears to be declining in some parts of its range, such that it is locally threatened in its postulated focal range but has, conversely, expanded into new areas as forest has been felled and converted to pasture. The main predator in Tasmania is the cat.

significance to humans

The eastern barred bandicoot was eaten by aboriginal Australians. It is a minor annoyance to landholders in suburban areas due to foraging in lawns.

Northern brown bandicoot

Isoodon macrourus

subfamily

Peramelinae

taxonomy

Perameles macroura (Gould, 1842), Port Essington, Northern Territory, Australia.

other common names

English: Brindled bandicoot, large northern bandicoot; German: Grosse Kurznasenbeutler.

physical characteristics

Head and body length is 11.8–18.5 in (300–470 mm); weight is 17.6–109.3 oz (500–3,100 g). Speckled brown-black fur, lighter on belly.

distribution

Western Australia, Northern Territory, Queensland, in which state it is coastal, New South Wales south to near Sydney, Australia. Also in southern New Guinea.

habitat

Areas of low ground cover including tall grass and low shrubs, irrespective of tree cover. Grassland, woodland, open forest, rarely in closed forest. Gardens in settled areas. In New Guinea, found in grasslands and grassy savanna woodlands as well as gardens.

behavior

Nocturnal, solitary except when courting or mating or females with young. Males are aggressive and use a gland behind the ear to scent-mark territory.

feeding ecology and diet

Omnivorous. Northern brown bandicoots mainly eat insects, earthworms, and other invertebrates, but also include berries, seeds, and plant fiber such as sugar cane in their diet. Food is obtained by digging conical pits with the strong forefeet after detection by smell.

reproductive biology

In the southern part of its range, breeding occurs from late winter to summer; further north breeding takes place year-round. There are eight nipples and litter size ranges from one to seven, usually two to four. The gestation period is 12.5 days and young are weaned at about two months. A chorioallantoic placenta is formed and retained after parturition. Northern brown bandicoots are polyestrous and promiscuous. Growth is rapid and sexual maturity is reached well before physical maturity. The nest is a heap of ground litter covering a shallow depression, with entrances at both ends. Hollow logs are also used.

conservation status

Overall, common to abundant, but has suffered local extinction due to altered habitat. Still present in urban areas in New South Wales, Queensland, and New Guinea, but such populations are at risk.

significance to humans

The northern brown bandicoot was eaten by Aboriginal Australians and is still hunted as food in New Guinea. It is considered a minor annoyance due to digging in suburban lawns.

Pig-footed bandicoot

Chaeropus ecaudatus

subfamily

Peramelinae

taxonomy

Perameles ecaudatus (Ogilby, 1838), Murray River, New South Wales, Australia.

other common names

German: Schweinfussnasenbeutler.

physical characteristics

Head and body length was 9.1–10.2 in (230–260 mm); weight was about 7 oz (200 g); comparatively long tail, despite its vernacular, bearing a terminal crest. Forefeet digits reduced to give appearance of pig's feet or deer hooves.

distribution

Formerly found across much of arid Australia, including much of inland Western Australia and South Australia, the southern half of the Northern Territory and marginally in western Queensland, New South Wales, and northwestern Victoria.

habitat

In the central deserts, occurred on sand dunes and sand plains with hummock grassland and tussock grass, sometimes with a

mulga (Acacia) overstory. In the east, occupied grassy plains and open woodland with a grass and shrub understory.

behavior

Nocturnal and presumably solitary except during mating and females with young. Gait compared with "a broken-down hack in a canter," but capable of explosive speed and agile leaps when disturbed.

feeding ecology and diet

The species' tooth structure and gut anatomy suggest that it was more herbivorous than other bandicoots. This notion is supported by observations of feeding by captives, and stomach content analysis. Termites, ants, and other insects were also eaten.

reproductive biology

Breeding is speculated to have been in May and June. The pouch has eight nipples, but litters seem to have been one to two. Probably promiscuous.

conservation status

Extinct. Last specimen collected in 1901 but Aboriginal testimony indicates that it probably survived in parts of its range until the 1950s.

significance to humans

None, although probably eaten by indigenous Australians.

Raffray's bandicoot

Peroryctes raffrayana

taxonomy

Perameles raffrayana (Milne-Edwards, 1878), Vogelkop, Irian Jaya, Indonesia.

other common names

English: Long-legged bandicoot; German: Grossen Neuguineanasenbeutler; Spanish: Bandicut de Raffray.

physical characteristics

Head and body length is 10.8–14.6 in (275–372 mm); weight is 22.9–35.3 oz (650–1,000 g). Unpatterned dark brown fur on back and on the long tail. Short, rounded ears.

distribution

Widespread in New Guinea, absent only from the woodlands and savanna of the south. Altitudinal range from about 130 to 13,120 ft (40 to 4,000 m), but most common at about 3,940 ft (1,200 m) along the central cordillera.

habitat

Lowland rainforest, lowland hill and mid-mountain oak forest, Nothofagus (beech) forest, mixed forest, and sub-alpine shrubs. Prefers undisturbed forest.

behavior

Not known, but probably nocturnal.

feeding ecology and diet

These bandicoots are reported to eat fruit, particularly that of figs.

reproductive biology

Females with pouch young have been captured between March and December, and the species may breed throughout the year. Litter size is one to two. It is possible that it nests communally, but that is not confirmed. One individual lived over three years in captivity. Mating is probably promiscuous.

conservation status

Common.

significance to humans

Eaten by indigenous people.

Rufous spiny bandicoot

Echymipera rufescens

taxonomy

Perameles rufescens (Peters and Doria, 1875), Kei Islands, Indonesia.

other common names

English: Long-nosed echymipera, spiny bandicoot, rufescent bandicoot; German: Dickkopf-Stachelnasenbeutler; Spanish; Echimipera Narizona.

physical characteristics

Head and body length is 11.8–16.1 in (300–410 mm); weight is 17.6–70.5 oz (500–2,000 g). Very elongate snout. Red-brown-black coarse, spiny dorsal fur, white ventrally. Short, almost naked black tail.

distribution

Subspecies E. r. rufescens is found in the lowlands of northern, eastern, and southern New Guinea, Aru Islands, and Kei Islands. E. r. australis is confined to Cape York, Australia.

habitat

In New Guinea is found only below 3,940 ft (1,200 m). Prefers rainforest but tolerates disturbed areas and grasslands. Australian subspecies occurs in closed forest including mesophyll vine forest, notophyll vine forest, and gallery forest, but also is found in eucalypt grassy woodland, coastal closed heath, and low layered open forest.

behavior

Nocturnal. Possibly uses burrows rather than nests for daytime shelter, at least in New Guinea.

feeding ecology and diet

Omnivorous, although prefers to eat insects.

reproductive biology

In New Guinea, pouch young have been recorded between March and October, but in Australia breeding may be more seasonal, with an estrus in the dry season. Possibly has lower fertility than seen in other species of the genus. Litter sizes from one to three are reported. Probably promiscuous.

conservation status

Generally uncommon, but may be locally abundant in New Guinea; common in its limited range on Cape York, Australia.

significance to humans

Eaten by indigenous humans in New Guinea.

Resources

Books

Flannery, T. Mammals of New Guinea. Sydney: Reed Books and Australian Museum, 1995.

Flannery, T. Mammals of the South-West Pacific and Moluccan Islands. Sydney: Reed Books and Australian Museum, 1995.

Gordon, G., and A. J. Hulbert. "Peramelidae." In Vol. 1B of Fauna of Australia: Mammalia, edited by D. W. Walton and B. J. Richardson. Canberra: Australian Government Publishing Service, 1989.

Jones, F. W. The Mammals of South Australia. Part 2. The Bandicoots and the Herbivorous Marsupials. Adelaide: Government Printer, 1924.

Mahoney, J. A., and W. D. L. Ride. "Peramelidae." In Zoological Catalogue of Australia 5, Mammalia. Canberra: Australian Government Publishing Service, 1988.

Menzies, J. A Handbook of New Guinea Marsupials and Monotremes. Madang, Papua, New Guinea: Kristen Press, Inc., 1991.

Seebeck, J. H., P. R. Brown, R. L. Wallis, and C. M. Kemper, eds. Bandicoots and Bilbies. Chipping Norton, Australia: Surrey Beatty & Sons, 1990.

Strahan, R., ed. The Mammals of Australia. Sydney: Reed Books and Australian Museum, 1995.

Stodart, E. "Breeding and Behaviour of Australian Bandicoots." In The Biology of Marsupials, edited by B. Stonehouse and D. Gilmore. London: Macmillan, 1977.

Periodicals

Freedman, L. "Skull and Tooth Variation in the Genus Perameles. Part 1. Anatomical Features." Records of the Australian Museum 27 (1967): 147.

Gordon, G., and B. C. Lawrie. "The Rufescent Bandicoot, Echymipera rufescens (Peters and Doria), on Cape York Peninsula." Australian Wildlife Research 5 (1977): 41.

Heinsohn, G. E. "Ecology and Reproduction of the Tasmanian Bandicoots (Perameles gunni and Isoodon obesulus)." University of California Publications in Zoology 80 (1966): 1.

Seebeck, John H. "Perameles gunnii." Mammalian Species Account No. 654.

John H. Seebeck, BSc, MSc, FAMS