Peramelemorphia

 New Guinea spiny bandicoot Echymipera kalubu, photographed by Michael Pennay.

Belongs within: Australidelphia.

The Peramelemorphia includes the bandicoots, a group of long-snouted Australian marsupials. Most species are insectivorous or carnivorous, though the now-extinct pig-footed bandicoot Chaeropus ecaudatus may have been a grazer (Long et al. 2002).

The shrinking world of bandicoots
Published 26 August 2017

A bandicoot is a very disagreeable animal to clean, therefore it should be done as soon after killing as possible, and then the flesh can be left in strong vinegar and water for a few hours before dressing. Sweet potatoes and onion make a good stuffing for bandicoot, which is good either boiled or baked.–Mrs Lance Rawson, Australian Enquiry Book of Household and General Information.

Golden bandicoots Isoodon auratus barrowensis, copyright Kathie Atkinson.

Back when I used to work on Barrow Island in the north-west of Australia, one of the more noticeable animals to be seen around the place was the golden bandicoot Isoodon auratus. In the evenings, the place seemed to absolutely swarm with them. About the size of a guinea pig, with no tails to speak of (bandicoots are actually born with fairly long tails but tend to lose them in the course of their quite vicious fights with one another; few if any individuals reach maturity with their tails intact), there was no question about their qualifications when it came to cuteness.

Bandicoots are a group of twenty-odd species of marsupial found in Australia and New Guinea (one species, the Seram bandicoot Rhynchomeles prattorum, was described from montane forest on the Indonesian island of Seram to the west of New Guinea). Most are primarily insectivorous, but they also eat varying amounts of small vertebrates and plant matter such as bulbs and fruit. The largest bandicoot, the giant bandicoot Peroryctes broadbenti, has been recorded to reach close to five kilograms in weight. The smallest, the Papuan bandicoot Microperoryctes papuensis, weighs less than 200 grams. I suspect many people in Australia assume that the name ‘bandicoot’ comes from one of the the Aboriginal languages, but it is in fact Indian (specifically Telugu) in origin. The original bandicoot Bandicota indica is a large rat that is widespread in southern Asia and Australian bandicoots were named for their resemblance to this species. Personally, I have maintained in the past that Australian bandicoots look more like rats than rats do: with their relatively long snouts, bandicoots bear a distinct resemblance to the sort of cartoon figure that comes to most people’s minds when they hear the word ‘rat’.

Bandicoots are highly distinctive from all other marsupials in appearance. Their hind legs are noticeably longer than their forelegs and more or less specialised for cursorial locomotion (especially so in one example that I’ll get to shortly). The fourth and fifth toes of the hind foot are much larger than the other three; the first toe in particular is reduced to a non-functional stub. The second and third toes of the hind foot, as in diprotodontian marsupials such as kangaroos and possums, are externally joined together with the two claws at the end forming a comb that is used in grooming more than in locomotion. The fore feet, in contrast, are mostly functionally three-fingered (with the first and fifth fingers reduced) and adapted for digging with the claws large and flat.

Many bandicoots are rapid reproducers with their gestation periods among the shortest of any mammal, less than two weeks between fertilisation and birth. Bandicoots also have the most developed placentas of any marsupial group (yes, most marsupials do have a placenta, albeit a much simpler one than found in placental mammals); it is presumably because of this that, despite their short gestation, bandicoot young are born at a more advanced stage of development than those of some other marsupials. When the young are born, they initially remain attached to their mother via the umbilical cord; this latter does not become detached and the placenta ejected until after the joey is firmly attached to a teat in the rearward-opening pouch. The young remain in the pouch for about two months and grow rapidly; they may reach full sexual maturity at the age of only three months. As a result, bandicoot populations may increase rapidly if conditions permit.

Greater bilby Macrotis lagotis, copyright Bernard Dupont.

In terms of classification, there is a general consensus that Recent bandicoots can be divided between four groups though there has been some disagreement about exactly these groups are interrelated (and hence exactly how they should be ranked). The most diverse, but probably also the least studied, group of modern bandicoots are the rainforest bandicoots of the Peroryctidae or Peroryctinae. These are about a dozen species found mostly in New Guinea with the aforementioned Rhynchomeles prattorum on Seram and the the long-nosed spiny bandicoot Echymipera rufescens extending its range to the northern tip of Queensland. Most of continental Australia is home to the dry-country bandicoots of the Peramelidae sensu stricto or Peramelinae, of which there are six Recent species (one of these, the northern brown bandicoot Isoodon macrourus, is also found in southern New Guinea). Peramelids tend to have shorter snouts and flatter skulls than peroryctids. The other two groups are both very small and also native to arid regions of Australia. Two Recent species are known of the genus Macrotis, the bilbies, though one of these is extinct and the other is endangered. Bilbies are larger than most other bandicoots, with long ears (hence their alternative vernacular name of ‘rabbit-bandicoots’) and a long, silky-haired tail.

Gerard Krefft’s 1857 illustration of the pig-footed bandicoot Chaeropus ecaudatus, from here.

The final representative of the Recent bandicoots is unquestionably the weirdest of them all. Unfortunately, it is also now extinct, last recorded some time about the middle of the 20th Century, a fact that cannot be called anything less than a fucking tragedy. The pig-footed bandicoot Chaeropus ecaudatus was the most cursorial of all bandicoots. Its forelegs, rather than being adapted for digging as in other bandicoots, had only two functional toes on which the claws were modified into hooves. The hind legs went a step further and had only a single functional toe (raising the question of how this animal groomed itself without the aforementioned claw-comb of other bandicoots Edit: That was a bit of a blonde moment; a second look at the Krefft illustration above shows that the comb is definitely there). The most extensive observations of its habits seem to have been made by Gerard Krefft (1866) who kept a pair alive for about six weeks in 1857 on a trip to the Murray-Darling region before killing them to provide specimens because, you know, 19th-Century naturalist. Krefft recorded that his bandicoots subsisted primarily on plant foods such as lettuce, grass and roots, refusing all meat offered to them (Krefft also refers to providing grasshoppers for them but his account is unclear about whether they were ever eaten). A herbivorous diet was also indicated by the animals’ droppings, which where dry and similar to a sheep’s. The bandicoots constructed a covered nest from grass and leaves in the tin enclosure in which Krefft kept them in which they sheltered during the day, only becoming active after nightfall. Krefft notes that he acquired “about eight” specimens of pig-footed bandicoot during his six-month camp, admitting that some met a stickier end than others: “They are very good eating, and I am sorry to confess that my appetite more than once over-ruled my love for science; but 24 hours upon “pig-face” (mesembryanthemum) will dampen the ardour of any naturalist“. Krefft also noted that several of the specimens found were female, and that despite being provided with eight teats the females never carried more than two joeys. A particularly interesting detail was that the fourth toe of the joeys’ fore foot, rather than being reduced as in the adults, remained large so that the feet resembled those of other bandicoots. Presumably this was so that the fore-claws could still be used to allow the newborn joeys to climb from the birth canal to the pouch.

Krefft also noted that the pig-footed bandicoot was already declining in abundance, blaming its increased rarity on competition with introduced grazing livestock. Sadly, changing habitats and introduced predators have caused other bandicoot species to also become endangered since Krefft’s time. Please, don’t let them go the way of the pig-footed bandicoot.

Systematics of Peramelemorphia

Characters (from Long et al. 2002): Lower molars with very large, un-notched anterior cingulid; upper molars with large adjacent stylar cusps C and D, and buccally shifted centrocrista. Hind feet syndactylous (at least in Perameloidea), with second and third toes reduced and joined by a web of skin).

Peramelemorphia [Peramelia, Perameliformes, Peramelina]
| i. s.: Numbigilga [Numbigilgidae]BA12
| `--N. ernielundeliusi Beck et al. 2008BA12
|--Yarala Muirhead & Filan 1995 [Yaralidae, Yaraloidea]LA02
| |--*Y. burchfieldi Muirhead & Filan 1995LA02
| `--Y. kida Schwartz 2006BA12
`--Perameloidea [Peramelidae]LA02
| i. s.: Thylacis obesulus (Shaw & Nodder 1797)S57 [=Perameles obesulusS57; incl. P. macroura Gould 1842S57, G42]
|--+--Chaeropus ecaudatusFS15
| `--ThylacomyidaeLA02
| |--Ischnodon Stirton 1955LA02
| | `--*I. australis Stirton 1955LA02
| |--ThylacomysWJ26 [=Thalacomys Blyth 1840W00]
| | |--‘Thalacomys’ lagotisW00
| | `--T. nigripesWJ26
| `--MacrotisFS15
| |--M. lagotis (Reid 1837)GM12
| | |--M. l. lagotisD81
| | `--M. l. grandisD81
| `--M. leucuraCB-E04
`--+--+--IsoodonFS15
| | |--I. macrourus (Gould 1842)CB-E04, K92
| | `--+--I. auratus (Ramsay 1887)CB-E04, GM12
| | `--I. obesulus (Shaw 1797)CB-E04, BD-D09
| `--Perameles Geoffroy 1804CB-E04, P02
| | i. s.: P. allinghamensis Archer 1976LA02
| | P. fasciataD81
| | P. harveyi Waterhouse 1842W42
| | P. lagotisG41
| | P. myosuraD81
| | P. wombeyensis Broom 1896F71
| |--P. bougainville Quoy & Gaimard 1924CB-E04, P02
| |--+--P. bowensis Muirhead, Dawson & Archer 1997P02, LA02
| | `--P. sobbei Price 2002P02
| `--+--P. gunnii Gray 1838CB-E04, P02
| `--+--*P. nasuta Geoffroy 1804LA02, CB-E04, LA02
| `--P. eremiana Spencer 1897FS15, P02
`--PeroryctidaeLA02
|--PeroryctesFS15
| |--P. broadbentiCB-E04
| |--P. longicaudaA82
| |--P. raffrayana (Milne-Edwards 1878)APB99
| `--P. tedfordiBA12
`--+--Microperoryctes Stein 1932FS15, SM93
| |--M. apliniFS15
| |--M. longicauda (Peters & Doria 1875)APB99
| | |--M. l. longicaudaAPB99
| | |--M. l. dorsalisAPB99
| | `--M. l. ornatusAPB99
| |--M. murinaCB-E04
| `--M. papuensisCB-E04
`--Echymipera Lesson 1842FS15, LA02
| i. s.: E. doreyanaTL70
|--+--E. claraFS15
| `--E. echinista Menzies 1990FS15, APB99
`--+--Rhynchomeles prattorumFS15
`--+--E. rufescens (Peters & Doria 1875)FS15, APB99
| |--E. r. rufescensR64
| `--E. r. australis Tate 1948R64
`--+--*E. kalubu (Lesson 1828)R64, FS15, R64 [=Perameles kalubuR64]
`--E. davidiFS15

*Type species of generic name indicated

References

[APB99] Aplin, K. P., J. M. Pasveer & W. E. Boles. 1999. Late Quaternary vertebrates from the Bird’s Head Peninsula, Irian Jaya, Indonesia, including descriptions of two previously unknown marsupial species. Records of the Western Australian Museum Supplement 57: 351–387.

[A82] Archer, M. 1982. A review of Miocene thylacinids (Thylacinidae, Marsupialia), the phylogenetic position of the Thylacinidae and the problem of apriorisms in character analysis. In: Archer, M. (ed.) Carnivorous Marsupials vol. 2 pp. 445–476. Royal Zoological Society of New South Wales.

[BD-D09] Beveridge, I. & M.-C. Durette-Desset. 2009. A new species of the nematode genus Copemania (Nematoda: Trichostrongylida), parasitic in the western quoll, Dasyurus geoffroii and short-nosed bandicoot, Isoodon obesulus from south-western Western Australia. Records of the Western Australian Museum 25 (3): 345–349.

[BA12] Black, K. H., M. Archer, S. J. Hand & H. Godthelp. 2012. The rise of Australian marsupials: a synopsis of biostratigraphic, phylogenetic, palaeoecologic and palaeobiogeographic understanding. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 983–1078. Springer.

[CB-E04] Cardillo, M., O. R. P. Bininda-Emonds, E. Boakes & A. Purvis. 2004. A species-level phylogenetic supertree of marsupials. Journal of Zoology 264: 11–31.

[D81] Day, D. 1981. The Doomsday Book of Animals: A unique natural history of three hundred vanished species. Ebury Press: London.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GM12] Gibson, L. A., & N. L. McKenzie. 2012. Occurrence of non-volant mammals on islands along the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 15–40.

Gordon, G., & A. J. Hulbert. 1989. Peramelidae. In: Fauna of Australia vol. 1B. Mammalia. Australian Biological Resources Study: Canberra.

[G42] Gould, J. 1842. Characters of a new species of Perameles, and a new species of Dasyurus. Proceedings of the Zoological Society of London 10: 41–42.

[G41] Gulliver, G. 1841. On the blood-corpuscles of marsupial animals. Proceedings of the Zoological Society of London 9: 49–51.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i–vi, 1–154.

Krefft, G. 1866. On the vertebrated animals of the lower Murray and Darling, their habits, economy, and geographical distribution. Transactions of the Philosophical Society of New South Wales 1862–1865: 1–33.

[LA02] Long, J., M. Archer, T. Flannery & S. Hand. 2002. Prehistoric Mammals of Australia and New Guinea: One Hundred Million Years of Evolution. University of New South Wales Press: Sydney.

[P02] Price, G. J. 2002. Perameles sobbei sp. nov. (Marsupialia, Peramelidae), a Pleistocene bandicoot from the Darling Downs, south-eastern Queensland. Memoirs of the Queensland Museum 48: 193–197.

[R64] Ride, W. D. L. 1964. A list of mammals described from Australia between the years 1933 and 1963 (comprising newly proposed names and additions to the Australian faunal list). Australian Mammal Society Bulletin 7 (Suppl.): 1–15.

[S57] Sandars, D. F. 1957. Redescription of some cestodes from marsupials. II.—Davaineidae, Hymenolepididae and Anoplocephalidae. Annals of Tropical Medicine and Parasitology 51 (3): 330–339.

[SM93] Stucky, R. K., & M. C. McKenna. 1993. Mammalia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 739–771. Chapman & Hall: London.

[TL70] Turnbull, W. D., & E. L. Lundelius Jr. 1970. The Hamilton fauna: a Late Pliocene mammalian fauna from the Grange Burn, Victoria, Australia. Fieldiana Geology 19: 1–163.

[W00] Waite, E. R. 1900. The generic name Thylacomys. Annals and Magazine of Natural History, series 7, 5: 222–223.

[W42] Waterhouse, G. R. 1842. On two new species of marsupial animals from South Australia. Proceedings of the Zoological Society of London 10: 47–48.

[WJ26] Wood Jones, F. 1926. The R. M. Johnston Memorial Lecture, 1925. The mammalian toilet and some considerations arising from it. Papers and Proceedings of the Royal Society of Tasmania 1925: 14–62.

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