Chaeropus, known as the pig-footed bandicoots, is a genus of small marsupials that became extinct during the 20th century. They were the only members of the family Chaeropodidae in order Peramelemorphia (bandicoots and bilbies), with unusually thin legs, yet were able to move rapidly. Two recognised species[3] inhabited dense vegetation on the arid and semiarid plains of Australia. The genus' distribution range was later reduced to an inland desert region, where it was last recorded in the 1950s; it is now presumed extinct.
The genus was proposed by William Ogilby in a presentation to the Linnean Society of London of a new species tentatively assigned to a genus of bandicoots, the long-nosed Perameles, and was forwarded to John Gould, then at Sydney, for a more detailed examination. Ogilby submitted a drawing by collector Major Mitchell, who also supplied extensive remarks on the animal's form and habits, and identified the unusual pig-like toes of the forelimbs as the basis for a new genus. The collection of the specimen was made at the interior of New South Wales by Mitchell, on the banks of the Murray River.[2]
The genus name Chaeropus combines terms from Ancient Greek for "pig" and "foot", a reference to the unique characteristic of the forelimbs noted by the describing author. The specific epithet ecaudatus refers to the absence of a tail, which the single specimen happened to be missing, leading to the unfortunate suggestion by the name that the marsupial did not possess them.[4]
The recognised synonymy of the genus was published by Theodore S. Palmer in 1904.[4] A nomenclatural synonym, Chœropus [Choeropus] Waterhouse, G.R. 1841, was published several years after Ogilby as an unjustified emendation;[5][6] Waterhouse gives the spelling proposed by Ogilby in the same work.[7]Oldfield Thomas noted the inappropriate epithet ecaudatus in 1888, substituting the name Chœropus castanotis proposed by John Edward Gray as the type of the genus,[8] but this was suppressed as a synonym by Palmer.
The vernacular description, pig-footed bandicoot, was given by Ogilby in his first description, a name ascribed by Mitchell's party.[2] Europeans settlers reported the species as resembling "small antelopes",[9] a simile that was reported by Bernard Woodward as persisting until their disappearance around 1900. The names recorded from the Noongar language are boda, woda, and boodal.[10] According to Indigenous Australian trackers, the pig-footed bandicoot was known as landwang and tubaija in their culture.[11][12]
Classification
This genus was previously placed in the familyPeramelidae, along with the bilbies, as the subfamily Chaeropodinae by McKenna and Bell (1997). However, its form is quite distinct, and recent molecular evidence supports this distinction. It is believed to be the sister group of the rest of the Peramelemorphia, and has been assigned to its own family, the Chaeropodidae.[1] The divergence of Chaeropus from other members of the Peramelemorphia is estimated to be in the Mid-Late Oligocene, around 26.7 Mya (21.9–31.3 confidence interval).[13]
Until 2019, both species were grouped under C. ecaudatus as the pig-footed bandicoot; however, a 2019 study split the genus into two species - the northern pig-footed bandicoot (C. yirratji) and the southern pig-footed bandicoot (C. ecaudatus).[14] In 2016, a fossil species C. baynesi was described from Late Pliocene to Early Pleistocene (2.47–2.92 Mya) of south-west New South Wales.[15]
The arrangement within the Marsupialia, with the treatment proposed in the 2019 revision, may be summarised as:
A genus of Peramelemorphia order of marsupials, allied by a monophyletic family, and regarded as exceptional in their morphology.[6] The direct evidence of the two species is limited to 29 specimens remaining at museums in Australia and overseas.[16] Their superficial appearance resembled the native bandicoots or rat kangaroos, although very small and dainty, and comparable to the size of a young rabbit or kitten.[17]
The feet of forelimbs resemble those of the genus Sus and the hind legs seen as similar to a horse.[17] The species had a body size of 23–26cm (9.1–10.2in) and a 10–15cm (3.9–5.9in) tail.[18] In form, they were almost bilby-like on first sight, having long, slender limbs, large, pointed ears, and a long tail. On closer examination, however, it became apparent that the pig-footed bandicoots were very unusual for marsupials. The forefeet had two functional toes with hooves, similar to the cloven hoof of a pig or deer;[19] this is possibly due to juveniles being deposited in the pouch through external stalks, thus relieving them of using the forelimbs while as joeys.[20] The hind feet had an enlarged fourth toe with a heavy claw shaped like a tiny horse's hoof, with the other toes being vestigial; only the fused second and third toes were useful, and that not for locomotion, but grooming.
They had broad heads, and a long yet slender snout. Their fur was coarse and straight, but not spiny. In color, they varied from grizzled grey through fawn to orange-brown, and the belly and underparts were white with the fur on the ears being of chestnut color.
The genus had five pairs of upper and three pairs of lower incisor teeth; tooth shape differed between the two species. The females of the genus had eight nipples and the opening of the pouch was faced backwards, not forwards as is the case with kangaroos.[21][22]
Depictions of the species include a lithograph by Henry C. Richter in John Gould's Mammals of Australia, published in the mid-19th century. An unpublished illustration by Richter, under the direction of Gould, was discovered in the archives of Knowsley Hall, at one time home to a great patron of natural history, the Earl of Derby.[23] The species C. ecaudatus was selected by Peter Schouten and Tim Flannery for a series of paintings illustrating the known information on the species' appearance and habits, published in a book surveying the modern extinctions of animals.[17] Another reconstruction of the genus was produced by Schouten to illustrate the species C. yirratji.[16]
Distribution and habitat
The two species were native to western New South Wales and Victoria, the southern part of the Northern Territory and South Australia and Western Australia.[21]Chaeropus ecaudatus populated semiarid southern regions of southern Australia, extending to southwest Australia, while C. yirratji populated sandy environments extending from Western Australia to the deserts in central Australia.[24] They inhabited a wide range of habitat types, from grassy woodland and grassland plains to the spinifex country and arid flats of central Australia. Despite its wide range, the genus had a sparse distribution and was never abundant.[17]
The number of complete specimens found in museum collections is 29.[16] Historical records of collections in central and western Australia, where it was sometimes locally common, begin with a specimen obtained at the Peron Peninsula during the Uranie expedition led by Louis de Freycinet in 1818. John Gilbert recorded his observations at dense stands of Casuarina seedlings (Allocasuarina species) at the interior of the southwest, beyond Northam, but did not appear in the Avon valley, inland from the new colony at Perth, at new settlements near Toodyay, York, and the Wongan Hills area. On his second collecting expedition in the southwest he recorded the species at King George Sound, on the southern coast of Western Australia.[10] The species was not recorded in the lists produced by the Austin Expedition of 1854, although W. A. Sanford was informed of a frequently seen animal, described as the "chestnut-eared hog's-foot", which Ludwig Glauert suggests was too common to merit notice in Austin's published report. Despite actively searching and collecting information on this animal, the species was not recorded by John Tunney, George Masters, or Guy Shortridge in their extensive surveys of western regions during the subsequent decades.[25] The last record of Chaeropus in the Southwest Australian region was in 1898, at Youndegin, consistent with the disappearance of species in a critical weight range that succumbed to Australia's mammalian faunal collapse (1875–1925).[26][10]
Archaeological evidence of the genus shows a greater range during Holocene and the earlier Pleistocene epoch. Specimens were uncovered in the first examination of fossil deposits at Mammoth Cave (Western Australia), at the beginning of the twentieth century, and great extension of the known range to eastern Queensland at two sites examined a century later.[27] The appearance in the fossil fauna of eastern Queensland sites demonstrates the replacement of rainforest communities with the open grassy habitat favoured by this genus during the later Pleistocene.[28] A survey of mammal fauna in the northeast of South Australia uncovered a fragment in an owl pellet buried beneath sediments.[29]
Explorer Samuel Albert White was shown the tracks of the animal in 1921 and gained information on its habitat, being found on stony tablelands or at sand dunes in central Australia, and was able to capture one with the assistance of a local Aboriginal woman and her dogs.[30] The increasing rarity of the animal had it included on a list of prohibited exports, either live or skin, issued by the federal government in 1921.[31]
Extinction
A 1902 illustration
According to Indigenous Australian oral tradition, pig-footed bandicoots were rare even before the arrival of Europeans on the continent,[22] and were in a serious decline even as it first came to scientific notice in the middle years of the 19th century. Two specimens of pig-footed bandicoots were obtained by local people in 1857 for Gerard Krefft, who accompanied the Blandowski Expedition. In trying to communicate the species he sought, Krefft showed a drawing to his collectors that showed the absence of a tail; the first specimens he was supplied were bandicoots with the tails removed. Despite the trouble taken in gaining living specimens, Krefft recorded his observations with an apology for eating one of them.[32][33] Only a handful of specimens was collected through the second part of the 19th century, mostly from northwestern Victoria, but also from arid country in South Australia, Western Australia, and the Northern Territory. By the start of the 20th century, they had become extinct in Victoria and the south-west of Western Australia. The last certain specimen was collected in 1901. By 1945, C. ecaudatus was extinct, having vanished from South Australia, and C. yirratji was reported to be limited to "a slight foothold in central Australia".[22] Nevertheless, Aboriginal people report that C. yirratji survived as late as the 1950s in the Gibson Desert and the Great Sandy Desert of Western Australia.[34]
The cause of the extinction remains uncertain; neither of the two most destructive introduced exterminator species, the fox and the rabbit, had yet arrived in south-west Western Australia when the pig-footed bandicoots disappeared from that area. Feral cats were already common, which may offer an explanation; perhaps more likely, the decline was caused by a double habitat change. Firstly, the end of many thousands of years of Aboriginal burning, which being confined to a patchwork of small areas at any one time, had ensured both fresh new growth in the recently burnt areas and adjacent older growth for shelter and as a base for recolonisation. Australia's Aboriginal population had declined by around 90% during the 19th century, largely because of the introduction of European diseases, and the remaining Aborigines were often no longer permitted to carry on their traditional land-management and hunting practices. Secondly, following on the heels of the near-extermination of the Aborigines, came the introduction of vast numbers of sheep and cattle, leading to significant changes in soil structure, plant growth, and food availability.[citation needed]
The species was included in historical modelling of a disease outbreak, a theorised epizootic that was the primary cause of mammalian declines, to which the populations of Chaeropus would seem to have been highly susceptible. The sudden demise of these marsupials was noted by Hal Colebatch, writing in 1929 that the disappearance was not the result of direct actions of settlers, but an unexplained consequence of a natural event. The apparent resilience of some mammals at Kellerberrin, which reappeared after a localised collapse, did not include this genus. When modelled as a primary factor in their demise, which has no discernible secondary factors, the population had no immunity to the hypothetical epizootic.[35]
An ethnographic survey of Aboriginal informants from central Australia in 1988 found the animal has not been seen in southern regions for around 70 years, but had persisted in the central desert regions until 30–50 years earlier.[36]
Few naturalists had the opportunity to observe and document the behaviour of the two species, with one of the few existing accounts suggesting that it moved "like a broken-down hack in a canter, apparently dragging the hind quarters after it".[37] This is contradicted by the Aboriginal people of central Australia, who knew it well and reported that if disturbed, it was capable of running with considerable speed by breaking into a smooth, galloping sprint.
Chaeropus species moved with a distinctive gait, exaggerated by their proportionally long and slender limbs that resemble a large grazing mammal like the African antelope.[10] Each forelimb had two functional toes and the rear limb ending in a hoof-like toe, with an apparent advantage when used to quickly evade a perceived threat.[17] They were solitary, nocturnal animals that would sleep in their shelter during the day and emerge in the evening to feed, using their keen sense of smell to find food.[18] Depending on the habitat, pig-footed bandicoots used a variety of shelters to hide from predators and for sleeping. John Gilbert found a nest site near Northam, Western Australia, that he described as greatly resembling those made by quenda and marl, but with more leaves.[10] In wooded areas and grasslands, these ranged from hollow logs to nests made out of grass, while in arid, treeless country, this animal dug short, straight burrows with a nest at the end.
From surviving eyewitness reports and analyses of gut contents, dentition, and gut structure of museum specimens, pig-footed bandicoots apparently were the most herbivorous of the peramelemorphs; analysis based on dental morphology and tooth wear are consistent with a predominately plant-based diet.[38] While captive specimens were fond of meat, and Aborigines reported that they ate grasshoppers, ants, and termites, the bulk of their diet was almost certainly leaves, roots, and grasses. In captivity, they drank "a good deal of water".[17]
Tim Flannery suggests that breeding occurred between May and June and that twins may have been the norm for this species. From the size of its pouch and comparison with other marsupials of this size, pig-footed bandicoots likely did not carry more than four young per litter.[18] Correspondence to Ludwig Glauert, reporting its disappearance some decades earlier, suggested that the animal was extremely delicate and would die if caught and mishandled.[26]
Related Research Articles
Macrotis is a genus of desert-dwelling marsupial omnivores known as bilbies or rabbit-bandicoots; they are members of the order Peramelemorphia. At the time of European colonisation of Australia, there were two species. The lesser bilby became extinct in the 1950s; the greater bilby survives but remains endangered. It is currently listed as a vulnerable species. The greater bilby is on average 55 cm (22 in) long, excluding the tail, which is usually around 29 cm (11 in) long. Its fur is usually grey or white; it has a long, pointy nose and very long ears, hence the reference of its nickname to rabbits.
The greater bilby, or simply bilby, is a long-eared, rabbit-like mammal native to Australia. It lives in burrows and is active at night, feeding on insects, fruit, or fungi. The bilby is a marsupial and carries its young in a pouch. Threats include habitat loss, disease, and introduced predators such as foxes. Formerly widespread, bilbies are now restricted to arid parts of northwestern and central Australia.
The order Peramelemorphia includes the bandicoots and bilbies. All members of the order are endemic to Australia-New Guinea and most have the characteristic bandicoot shape: a plump, arch-backed body with a long, delicately tapering snout, very large upright ears, relatively long, thin legs, and a thin tail. Their size varies from about 140 grams up to 4 kilograms, but most species are about one kilogram.
Bandicoots are a group of more than 20 species of small to medium-sized, terrestrial, largely nocturnal marsupial omnivores in the order Peramelemorphia. They are endemic to the Australia–New Guinea region, including the Bismarck Archipelago to the east and Seram and Halmahera to the west.
The desert bandicoot is an extinct bandicoot of the arid country in the centre of Australia.
The southern pig-footed bandicoot was a small species of herbivorous marsupial in the genus Chaeropus, the pig-footed bandicoots.
Australidelphia is the superorder that contains roughly three-quarters of all marsupials, including all those native to Australasia and a single species — the monito del monte — from South America. All other American marsupials are members of the Ameridelphia. Analysis of retrotransposon insertion sites in the nuclear DNA of a variety of marsupials has shown that the South American monito del monte's lineage is the most basal of the superorder.
The lesser bilby, also known as the yallara, the lesser rabbit-eared bandicoot or the white-tailed rabbit-eared bandicoot, was a rabbit-like marsupial. The species was first described by Oldfield Thomas as Peregale leucura in 1887 from a single specimen from a collection of mammals of the British Museum. Reaching the size of a young rabbit, this species lived in the deserts of Central Australia. Since the 1950s–1960s, it has been believed to be extinct.
Perameles is a genus of marsupials of the order Peramelemorphia. They are referred to as long-nosed bandicoots or barred bandicoots.
The short-nosed bandicoots are members of the order Peramelemorphia. These marsupials can be found across Australia, although their distribution can be patchy. Genetic evidence suggests that short-nosed bandicoots diverged from the related long-nosed species around eight million years ago, during the Miocene epoch, and underwent a rapid diversification around three million years ago, during the late Pliocene.
The marsupial family Peramelidae contains the extant bandicoots. They are found throughout Australia and New Guinea, with at least some species living in every available habitat, from rainforest to desert. Four fossil peramelids are described. One known extinct species of bandicoot, the pig-footed bandicoot, was so different from the other species, it was recently moved into its own family.
The southern brown bandicoot is a short-nosed bandicoot, a type of marsupial, found mostly in southern Australia. A subspecies in Western Australia was also known as the quenda in South Western Australia. This subspecies was elevated to species in 2018.
The northern pig-footed bandicoot was a small species of extinct herbivorous Australian marsupial in the genus Chaeropus, the pig-footed bandicoots. It has been believed to be extinct since the mid-20th century; the last confirmed observation was a specimen collected near Alice Springs in 1901, but reports from local Aborigines indicate that it may have survived in the Gibson and Great Sandy Deserts as late as the 1950s.
Crash bandicoot is an extinct bandicoot, known from fossils located at the Riversleigh World Heritage Area in northeast Australia.
Liyamayi dayi is a mammal species of the Thylacomyidae family known from fossils located at the Riversleigh World Heritage Area in northeast Australia. The discovery of the specimens was identified as deposited around fifteen million years ago, revising the earliest record of this peramelemorphian lineage from those of species that existed around ten million years later.
Perameles myosuros, the south-western barred bandicoot or marl, is a recently extinct species of bandicoot that was native to the southern parts of Western Australia. Initially described in 1841 it was later made a subspecies of the extant Perameles bougainville before being restored to species level in 2018, based on museum specimens. Of particular note was that the females were significantly larger than the males in P. myosuros, which is an unusual trait amongst mammals, whereas the sexes were the same size in P. bougainville.
↑ Louys, Julien; Cramb, Jonathan; Price, Gilbert J.; Stemmer, David; Brewer, Philippa; Brace, Selina; Miguez, Roberto Portela; Simões, Bruno F.; Travouillon, Kenny J. (2019-03-13). "Hidden in plain sight: reassessment of the pig-footed bandicoot, Chaeropus ecaudatus (Peramelemorphia, Chaeropodidae), with a description of a new species from central australia, and use of the fossil record to trace its past distribution". Zootaxa. 4566 (1): zootaxa.4566.1.1. doi:10.11646/zootaxa.4566.1.1. ISSN1175-5334. PMID31716448. S2CID92165477.
↑ Westerman, M.; Kear, B.P.; Aplin, K.; Meredith, R.W.; Emerling, C.; Springer, M.S. (January 2012). "Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences". Molecular Phylogenetics and Evolution. 62 (1): 97–108. doi:10.1016/j.ympev.2011.09.009. PMID22100729.
↑ Article in Journal of Mammalian Evolution 20(4) · December 2013 DOI: 10.1007/s10914-012-9220-3
↑ Cooper, W.J.; Steppan, S.J. (2010). "Developmental constraint on the evolution of marsupial forelimb morphology". Australian Journal of Zoology. 58 (1): 1. doi:10.1071/ZO09102. S2CID15693318.
↑ Fisher, C.T. (September 1988). "An Unpublished Drawing of the Pig-Footed Bandicoot by John Gould and H. C. Richter". Australian Zoologist. 24 (4): 205–209. doi:10.7882/AZ.1988.003.
↑ White, S.A. (29 July 1922). "Territory Flora and Fauna". The Register. Vol.LXXXVII, no.25, 456. Adelaide. p.9. Retrieved 27 July 2019– via National Library of Australia.
↑ "Australian Mammals". The Daily Telegraph. No.13292. New South Wales, Australia. 16 December 1921. p.4. Retrieved 27 July 2019– via National Library of Australia.
↑ Abbott, I. (December 2006). "Mammalian faunal collapse in Western Australia, 1875–1925: the hypothesised role of epizootic disease and a conceptual model of its origin, introduction, transmission, and spread". Australian Zoologist. 33 (4): 530–561. doi:10.7882/az.2006.024. ISSN0067-2238.
Waterhouse, G.R. (1841). Jardine, William (ed.). "Marsupialia, or pouched animals /". The Naturalist's Library. Conducted by Sir William Jardine, Bart. F.R.S.E., F.L.S., &c. &c. Edinburgh: W.H. Lizars. 11: 163.
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