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The order Peramelemorphia includes the bandicoots and bilbies; it equates approximately to the mainstream of marsupial omnivores All members of the order are endemic to the twin land masses of 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, or the weight of a half-grown kitten


  • 1 Phylogeny
    • 11 Placement within Marsupialia
    • 12 Relationships within Peramelemorphia
  • 2 Fossil record
  • 3 References


Placement within Marsupialia

The position of the Peramelemorphia within the marsupial family tree has long been puzzling and controversial There are two morphological features in the order that appear to show a clear evolutionary link with another marsupial group: the type of foot, and the teeth Unfortunately, these clear signposts point in opposite directions

All members of the order are polyprotodont have several pairs of lower front teeth—in the case of the Peramelemorphia, three pairs This suggests that they have evolved within Dasyuromorphia marsupial carnivores On the other hand, they also have an unusual feature in their feet: the second and third toes are fused together This condition is called syndactyly, and is characteristic of the Diprotodontia the order of marsupial herbivores that includes kangaroos, wombats, possums, and many others

Attempts to resolve this puzzle include the view that the bandicoot group evolved from the carnivores, retaining the polyprotodont dentition, and independently evolving a syndactyl hind foot; the contrary view that syndactyly is so unusual that it is unlikely to have evolved twice and therefore the bandicoot group must have evolved from a possum-like diprotodont creature, and re-evolved its extra teeth A third view suggests that the bandicoot group evolved from a primitive carnivore, developed the syndactylous hind foot as a specialisation for climbing, and the diprotodonts then split off and evolved the two-tooth jaw that gives them their name Recent molecular level investigations do not so far appear to have resolved the puzzle, but do strongly suggest that whatever the relationship of the bandicoot group to the other marsupial orders may be, it is a distant one

Relationships within Peramelemorphia

Recent molecular analyses have resulted in a phylogenetic reconstruction of the members of Peramelemorphia with quite strong support The most basal split separates Thylacomyidae Macrotis from all other bandicoots Probably the next to diverge was the recently extinct Chaeropodidae Chaeropus The remaining taxa comprise the Peramelidae, which divides into subfamilies Peramelinae Isoodon and Perameles and a clade in which the Echymiperinae Echymipera and Microperoryctes form a sister group to Peroryctinae Peroryctes:













Fossil record

Many specimens of modern peramelemorphian eg Perameles spp and Isoodon spp have been recovered in the fossil record from Pleistocene and Holocene fossil localities However, very few fossil species have been recovered to date The first species of fossil peramelemorphian was described by R A Stirton in 1955 The specimen Stirton described was a partial lower jaw from the Tirari Desert in Central Australia, Pliocene in age The lower jaw morphology suggested a relationship with bilbies Family Thylacomyidae, and was named Ischnodon australis

It was not until 1976 that Archer and Wade described the next fossil bandicoot A single upper molar was recovered from the Bluff Downs fossil site, Allingham Formation, in northern Queensland, also Pliocene in age The tooth was similar to that of species of Perameles, and was therefore named Perameles allinghamensis

In 1995, the first Miocene species was described from Riversleigh, and was named Yarala burchfieldi by Dr Jeannette Muirhead The species was represented by several upper and lower jaws, which were smaller than any living bandicoots and had a very primitive dentition A skull was later recovered in 2000, the first for any fossil peramelemorphian to date Features of the skull and dentition suggested that Yarala burchfieldi was distinct from other peramelemorphians, and for this reason, a new Superfamily Yaraloidea and Family Yaralidae were erected to classify this species

In 1997, Muirhead, Dawson and Archer described a new species of Perameles, Perameles bowensis, from teeth recovered from two Pliocene fossil localities, Bow and Wellington Caves The same species was later reported in 2000 from Chinchilla, Queensland by Mackness and colleagues In 2002, Price described a new species Perameles, Perameles sobbei, from the Darling Downs Pleistocene in age, south-eastern Queensland This species was represented by a lower jaw and a few isolated lower molars Additional material were later described in 2005 from the same site, including upper molars

A second species of Yarala, Yarala kida, was described in 2006 by Schwartz This species was recovered from Kangaroo Well, a late Oligocene site from the Northern Territory in Australia This species is thought to be even more primitive than Yarala burchfieldi

The second fossil skull of any fossil peramelemorphian was also recovered from Miocene sites of Riversleigh In fact, more than one skull of this new species was found and several lower and upper jaws, and was significantly different from any other bandicoot to erect a new genus, Galadi The species was named Galadi speciosus by Travouillon and colleagues It was short-snouted unlike modern bandicoots suggesting that it was more carnivorous than its omnivorous modern relatives Its relationship to other bandicoots is unclear, but it was likely to be less primitive than Yarala but more primitive than living bandicoots An additional three species of Galadi were later described in 2013 and named Galadi grandis, Galadi amplus and Galadi adversus

Gurovich et al 2013 described a new species of mouse-sized bandicoot from Riversleigh and from Kutjamarpu, Southern Australia The species, named Bulungu palara, is represented by a skull and several lower and upper jaws Two other species in this genus were also described from the Etadunna Formation in South Australia, Bulungu muirheadae which was the oldest fossil bandicoot recovered as of 2013 about 24 million years old, and Bulungu campbelli

The oldest modern bandicoot peramelid and the oldest bilby Thylacomyid were later discovered by Travouillon et al,2014 from Riversleigh World Heritage Area, from middle Miocene fossil deposits around 15 million years old The peramelid, Crash bandicoot, was named after the famous video game character and is only represented by a single upper jaw The bilby, Liyamayi dayinamed after geologist and philanthropist Robert Day, is only known from 3 teeth 2 upper molar, 1 lower molar

The first record of sexual dimorphism difference in size between males and females in a fossil bandicoot was reported from two new species from Riversleigh Travouillon et al 2014 Named Madju variae and Madju encorensis, they are closely related to modern bandicoots, but do not fall in any modern family, as did Galadi and Bulungu genus Instead they are classified as Perameloid, with all known Peremelemorphian, to the exclusion of Yaralids Madju variae is also unusual in preserving an ontogenetic series age series from pouch young to adult, the second of any fossil marsupial mammal in Australia The study of this ontogenetic series lead researchers to think that Madju variae developed slow than modern bandicoots, much more like a bilby, and therefore the rapid development of modern bandicoots must have evolved after the middle Miocene, when Australia started to become more arid


  1. ^ Aplin, KP, Helgen, KM, Lunde, DP, 2010 A review of Peroryctes broadbenti, the giant bandicoot of Papua New Guinea American Museum Novitates 3696, 1-41
  2. ^ Gordon, G, Hulbert, AJ, 1989 Peramelidae, In: Walton, DW Ed, Fauna of Australia Australian Government Publishing Service, Canberra, pp 603–624
  3. ^ Strahan, R 1995 Mammals of Australia Washington, DC: Smithsonian Institution Press
  4. ^ a b Meredith, Robert W; Westerman, Michael; Springer, Mark S 2008 "A timescale and phylogeny for "Bandicoots" Peramelemorphia: Marsupialia based on the sequences for five nuclear genes" Molecular Phylogenetics and Evolution 47 1: 1–20 doi:101016/jympev200801002 PMID 18328736 
  5. ^ Rich, T H, Archer, M, Hand, S J, Godthelp, H, Muirhead, J, Pledge, N S, Flannery, T F, Woodburne, M O, Case, J A, Teford, R H, Turnbull, W D, Lundelius, E Ljr, Rich, L S V, Whitelaw, M J, Kemp, A, & Rich, P V 1991 Australian Mesozoic and Tertiary terrestrial mammal localities, Appendix 1 Pp 1005-1058 in P Vickers-Rich, Monaghan J M, R F Baird & T H Rich eds, Vertebrate Palaeontology of Australia Pioneer Design Studio and Monash University Publications Committee, Melbourne
  6. ^ Stirton, R A 1955 Late Tertiary marsupials from South Australia Records of the South Australian Museum, 11, 247– 267
  7. ^ Archer, M & Wade, M 1976 Results of the Ray E Lemley expeditions, part 1: The Allingham Formation and a new Pliocene vertebrate fauna from northern Queensland Memoirs of the Queensland Museum, 17, 54–58
  8. ^ Muirhead, J & Filan, S L 1995 Yarala burchfieldi, a plesiomorphic bandicoot Marsupialia, Peramelemorphia from Oligo-Miocene deposits of Riversleigh, northwestern Queensland Journal of Paleontology, 691, 127-134
  9. ^ Muirhead, J 2000 Yaraloidea Marsupialia, Peramelemorphia, a new superfamily of marsupial and a description and analysis of the cranium of the Miocene Yarala burchfieldi Journal of Paleontology, 743, 512-523
  10. ^ Muirhead, J, Dawson, L & Archer, M 1997 Perameles bowensis, a new species of Perameles Peramelomorphia, Marsupialia from Pliocene faunas of Bow and Wellington caves, New South Wales Proceedings of the Linnean Society of New South Wales, 17, 163–174
  11. ^ Mackness, B S, Wroe, S, Muirhead, J, Wilkinson, C & Wilkinson, D 2000 First fossil bandicoot from the Pliocene Chinchilla Local Fauna Australian Mammalogy, 22, 133– 136
  12. ^ 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
  13. ^ Price, G J 2005 Fossil bandicoots Marsupialia, Peramelidae and environmental change during the Pleistocene on the Darling Downs, southeastern Queensland, Australia Journal of Systematic Palaeontology, 4, 347-356
  14. ^ Schwarz, L R S 2006 A new species of bandicoot from the Oligocene of northern Australia and implications of bandicoots for correlating Australian Tertiary mammal faunas Palaeontology, 49, 991–998
  15. ^ KJ Travouillon; Y Gurovich; RMD Beck; J Muirhead 2010 "An exceptionally well-preserved short-snouted bandicoot Marsupialia; Peramelemorphia from Riversleigh's Oligo-Miocene deposits, northwestern Queensland, Australia" Journal of Vertebrate Paleontology 30 5: 1528–1546 doi:101080/027246342010501463
  16. ^ K J Travouillon, Y Gurovich, M Archer, S J Hand and J Muirhead 2013 "The genus Galadi: three new bandicoots Marsupialia, Peramelemorphia from Riversleigh’s Miocene deposits, northwestern Queensland, Australia" Journal of Vertebrate Paleontology 33 1: 153–168 doi:101080/027246342012713416
  17. ^ Gurovich, Y, Travouillon, KJ, Beck, RMD, Muirhead, J, Archer, M, 2013 Biogeographical implications of a new mouse-sized fossil bandicoot Marsupialia: Peramelemorphia occupying a dasyurid-like ecological niche across Australia Journal of Systematic Palaeontology
  18. ^ Travouillon, KJ, Beck, RMD, Hand, SJ, Archer, M 2013 "The oldest fossil record of bandicoots Marsupialia; Peramelemorphia from the late Oligocene of Australia" Palaeontologia Electronica 16 2: 13A 52p CS1 maint: Multiple names: authors list link
  19. ^ Travouillon, KJ, Hand, S J, Archer, M, and Black, K H, 2014 Earliest modern bandicoot and bilby Marsupialia, Peramelidae and Thylacomyidae from the Miocene of the Riversleigh World Heritage Area, northwestern Queensland, Australia Journal of Vertebrate Paleontology 34:375-382
  20. ^ Travouillon, K J, Archer, M, Hand, S J and Muirhead, J, 2014 Sexually dimorphic bandicoots Marsupialia: Peramelemorphia from the Oligo-Miocene of Australia, first cranial ontogeny for fossil bandicoots and new species descriptions Journal of Mammalian Evolution doi:101007/s10914-014-9271-8
  • Groves, CP 2005 Wilson, DE; Reeder, DM, eds Mammal Species of the World: A Taxonomic and Geographic Reference 3rd ed Baltimore: Johns Hopkins University Press pp 38–42 ISBN 0-801-88221-4 OCLC 62265494 
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