Thylacoleo: Difference between revisions

Thylacoleo; Temporal range: late Pliocene—late Pleistocene
	Skeletal diagram of T. carnifex (top) and restored musculature based on living marsupials (bottom)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Infraclass:	Marsupialia
Order:	Diprotodontia
Family:	†Thylacoleonidae
Genus:	†Thylacoleo; Owen, 1859
Type species
	†Thylacoleo carnifex;
Species
	†T. carnifex; †T. crassidentatus; †T. hilli;

Browse history interactively

← Previous edit

Content deleted Content added

VisualWikitext

Inline

Latest revision as of 15:10, 6 December 2024

Thylacoleo ("pouch lion") is an extinct genus of carnivorous marsupials that lived in Australia from the late Pliocene to the Late Pleistocene (until around 40,000 years ago), often known as marsupial lions. They were the largest and last members of the family Thylacoleonidae, occupying the position of apex predator within Australian ecosystems. The largest and last species, Thylacoleo carnifex, approached the weight of a lioness. The estimated average weight for the species ranges from 101 to 130 kg (223 to 287 lb).^[1]

Taxonomy

The first Thylacoleo fossil findings, discovered by Thomas Mitchell were found in the 1830s in the Wellington Valley of New South Wales, though not recognised as such at the time. The generic holotype, consisting of broken teeth, jaws, and a skull was discovered by a pastoralist, William Avery, near Lake Colungolac from which the species Thylacoleo carnifex was described by Richard Owen.^[2] It was not until 1966 that the first nearly-complete skeleton was found. The only pieces missing were a foot and the tail. Currently, the Nullarbor Plain of West Australia remains to be the greatest finding site. These fossils now reside at the Australian Museum.^[3]^[4]

The genus was first published in 1859, erected to describe the type species Thylacoleo carnifex. The new taxon was established in examination of fossil specimens provided to Richard Owen. The familial alliance takes its name from this description, the so-called marsupial lions of Thylacoleonidae.

The colloquial name "marsupial lion" alludes to the genus name, which was named after its superficial resemblance to the placental lion and its ecological niche as a large predator.

Genus: Thylacoleo (Thylacopardus) – Australia's marsupial lions, that lived from about 2 million years ago, during the Late Pliocene Epoch and became extinct about 40,000 years ago, during the Late Pleistocene Epoch. Three species are known:

Thylacoleo carnifex The holotype cranium was collected from Lake Colongulac in 1843 by pastoralist William Adeney. A partial rostrum collected by Adeney in 1876 from the same locality would later be found to belong to the same individual.^[5] It was not until 1966 that the first nearly-complete skeleton was found.^[3]^[4]
Thylacoleo crassidentatus lived during the Pliocene, around 5 million years ago, and was about the size of a large dog. Its fossils have been found in southeastern Queensland.^[6]^[7]
Thylacoleo hilli lived during the Pliocene and was half the size of T. crassidentatus. It is the oldest member of the genus.^[8]

Fossils of other representatives of Thylacoleonidae, such as Microleo and Wakaleo, date back to the Late Oligocene Epoch, some 24 million years ago.^[9]

T. hilli was described by Neville Pledge in a study published in the records of the South Australia Museum in 1977. The holotype is a third premolar, discovered at a cave in Curramulka in South Australia, exhibiting the carnivorous characteristics of the genus and around half the size of T. carnifex. This tooth was collected by Alan Hill, a speleologist and founding member of the Cave Exploration Group of South Australia, while examining a site known as the "Town Cave" in 1956; the specific epithet hilli honours the collector of the first specimen.^[8] Material found amidst the fauna at Bow River in New South Wales, dated to the early Pliocene, was also referred to the species in 1982.^[10] A fragment of an incisor, unworn and only diagnosable to the genus, was located at a site in Curramulka, close to the Town Cave site, and referred to the species for the apparent correlation in size when compared to the better known T. carnifex.^[11]

The marsupial lion is classified in the order Diprotodontia along with many other well-known marsupials such as kangaroos, possums, and the koala. It is further classified in its own family, the Thylacoleonidae, of which three genera and 11 species are recognised, all extinct. The term marsupial lion (lower case) is often applied to other members of this family. Distinct possum-like characteristics led Thylacoleo to be regarded as members of Phalangeroidea for a few decades. Though a few authors continued to hint at phalangeroid affinities for thylacoleonids as recently as the 1990s, cranial and other characters have generally led to their inclusion within vombatiformes, and as stem-members of the wombat lineage.^[12] Marsupial lions and other ecologically and morphologically diverse vombatiforms were once represented by over 60 species of carnivorous, herbivorous, terrestrial and arboreal forms ranging in size from 3 kg to 2.5 tonnes. Only two families represented by four herbivorous species (koalas and three species of wombat) have survived into modern times and are considered the marsupial lion's closest living relatives.^[13]

Evolution

The ancestors of thylacoleonids are believed to have been herbivores, something unusual for carnivores. They are members of the Vombatiformes, an almost entirely herbivorous order of marsupials, the only extant representatives of which are koalas and wombats, as well as extinct members such as the diprotodontids and palorchestids.^[14] The group first appeared in the Late Oligocene. The earliest thylacoleonids like Microleo were small possum-like animals,^[15] with the group increasing in size during the Miocene, with representatives like the leopard-sized Wakaleo. The genus Thylacoleo first appeared during the Pliocene, and represented the only extant genus of the family from that time until the end of the Pleistocene. The youngest representative of Thylacoleo and the thylacoleonids, T. carnifex, is the largest known member of the family.^[14] The earliest thylacoleonids are thought to have been arboreal (tree dwelling) animals,^[16] while Thylacoleo is thought to be terrestrial with some climbing capabilities.^[17]

Description

T. carnifex is the largest carnivorous mammal known to have ever existed in Australia, and one of the largest metatherian carnivores known (comparable to Thylacosmilus and Borhyaena species, but smaller than Proborhyaena gigantea). Individuals ranged up to around 75 cm (30 in) high at the shoulder and about 150 cm (59 in) from head to tail. Measurements taken from a number of specimens show they averaged 101 to 130 kg (223 to 287 lb) in weight, although individuals as large as 124–160 kg (273–353 lb) might not have been uncommon, and the largest weight was of 128–164 kg (282–362 lb). This would make it comparable to female lions and female tigers in general size.^[17] Estimates of the size of T. carnifex based on dental remains are typically dubious, in contrast to estimates based on proximal limb bone circumference.^[18]

Skull

Like other thylacoleonids, Thylacoleo had blade-like third premolar teeth in the upper and lower jaws, that functioned as the carnassial teeth, with these teeth being present much further forwards in the jaw than in other mammals.^[19]^[20] Compared to earlier thyacoleonids, the third premolars were considerably enlarged.^[21] Thylacoleo also had a proportionally large pair of first incisors in the upper and lower jaws, which functioned analogously to other carnivores canine teeth.^[19]^[22] They also had true canines but they served little purpose as they were stubby and not very sharp.^[23] Compared to earlier thylacoleonids, the number of molar teeth was reduced.^[21]

Pound for pound, T. carnifex had the strongest bite of any mammal species, living or extinct; a T. carnifex weighing 101 kg (223 lb) had a bite comparable to that of a 250 kg African lion, and research suggests that Thylacoleo could hunt and take prey much larger than itself.^[20] Larger animals that were likely prey include Diprotodon spp. and giant kangaroos. It seems improbable that Thylacoleo could achieve as high a bite force as a modern-day lion; however, this might have been possible when taking into consideration the size of its brain and skull. Carnivores usually have rather large brains when compared to herbivorous marsupials, which lessens the amount of bone that can be devoted to enhancing bite force. Thylacoleo however, is thought to have had substantially stronger muscle attachments and therefore a smaller brain.^[24] Some later studies questioned the ability of the canine teeth to deliver a killing bite.^[25]

Using 3D modeling based on X-ray computed tomography scans, marsupial lions were found to be unable to use the prolonged, suffocating bite typical of living big cats. They instead had an extremely efficient and unique bite; the incisors would have been used to stab at and pierce the flesh of their prey while the more specialised carnassials crushed the windpipe, severed the spinal cord, and lacerated the major blood vessels such as the carotid artery and jugular vein. Compared to an African lion which may take 15 minutes to kill a large catch, the marsupial lion could kill a large animal in less than a minute.^{[dubious – discuss]} The skull was so specialized for big game that it was very inefficient at catching smaller animals, which possibly contributed to its extinction.^[26]^[27]

Postcranium

Thylacoleo had highly mobile and powerful forelimbs used to grapple prey, with each manus having a single very large retractable hooked claw set on large semi-opposable thumbs, which are suggested to have been used deal a killing blow.^[25]

The hind feet had four functional toes, the first digit being much reduced in size, but possessing a roughened pad similar to that of possums, which may have assisted with climbing. The discovery in 2005 of a specimen which included complete hind feet provided evidence that the marsupial lion exhibited syndactyly (fused second and third toes) like other diprotodonts.^[28]

Its strong forelimbs and retracting claws mean that Thylacoleo possibly climbed trees and perhaps carried carcasses to keep the kill for itself (similar to the leopard today).^[29] The climbing ability would have also helped them climb out of caves, which could therefore have been used as dens to rear their young.^[30] Specialised tail bones called chevrons strengthened the tail, likely allowing the animal to use it to prop itself up while rearing on its hind legs, which may have been done when climbing or attacking prey.^[30]

The lumbar region is relatively rigid and straight, and suggests that the lower back was relatively inflexible.^[30]

Ecology

When Thylacoleo was first described by Richard Owen, he considered it to be a carnivore, based on the morphology of its skull and teeth.^[2] However other anatomists, such as William Henry Flower disagreed. Flower was the first to place Thylacoleo with the Diprotodonts, noting its skull and teeth to be laid out more like those of the koala and the wombat, and suggested that it was more likely a herbivore. Owen did not disagree with Flower's placement of Thylacoleo with the Diprotodonts, but still maintained that it was a carnivore, despite its herbivorous ancestry.^[31] Owen found little support in his lifetime, despite the pointing out of Thylacoleo's retractable claws, something only found in mammalian carnivores,^[4] and its lack of any ability to chew plant material.^[31] In 1911, a study by Spencer and Walcott claimed that certain marks on the bones of megafauna had been made by Thylacoleo, but according to Horton (1979) they were not sufficiently rigorous, resulting in their arguments being strongly challenged by later scholars, such as Anderson (1929), and later Gill (1951, 1952, 1954), thereby leaving the issue unresolved.^[32]

Besides the most common hypothesis that it was an active predator, a variety of other theories existed in the late 19th to early 20th centuries as to the diet and feeding of Thylacoleo, with hypotheses of it being a scavenger filling the ecological niche of hyenas,^[33] being a specialist of crocodile eggs,^[29] or even a melon-eater.^[34] As late as 1954, doubts were still being raised as to whether it was actually a hypercarnivore.^[25]

In 1981, another paper was published arguing that certain cuts to bones of large marsupials had been caused by Thylacoleo. This paper by Horton and Wright was able to counter earlier arguments that such marks were the result of humans, largely by pointing out the presence of similar marks on the opposite side of many bones. They concluded that humans were extremely unlikely to have made the marks in question, but that if so "they had set out to produce only marks consistent with what Thylacoleo would produce".^[32] Since then, the academic consensus has emerged that Thylacoleo was a predator and a hypercarnivore.^[25]

The marsupial lion's limb proportions and muscle mass distribution indicate that, although it was a powerful animal, it was not a particularly fast runner. Paleontologists conjecture that it was an ambush predator, possibly using leaping.^[30] Incisions on bones of the extinct large kangaroo Macropus titan, and the general morphology of Thylacoleo suggests that it fed in a similar manner to modern cheetahs, by using their sharp teeth to slice open the ribcage of their prey, thereby accessing the internal organs. They may have killed by using their front claws as either stabbing weapons or as a way to grab their prey with strangulation or suffocation.^[32]

Like many predators, it was probably also an opportunistic scavenger, feeding on carrion and driving off less powerful predators from their kills. It also may have shared behaviours exhibited by recent diprotodont marsupials such as kangaroos, like digging shallow holes under trees to reduce body temperature during the day.^[35]

Trace fossils in the form of claw marks and bones from caves in Western Australia analyzed by Gavin Prideaux et al. indicate marsupial lions could also climb rock faces, and likely reared their young in such caves as a way of protecting them from potential predators.^[36]

Analysis of finds on the Nullabor Plain suggests that Thylacoleo carnifex inhabited open, arid environments similar to those found across much of Australia today.^[37]

A 1985 study suggested that Thylacoleo carnifex was an apex predator that primarily fed on large bodied prey, which may have included the large kangaroos Sthenurus, Procoptodon, Protemnodon, Macropus and Ostphranter as well as possibly the largest Australian marsupial, the rhinoceros-sized vombatoid Diprotodon.^[38]

Extinction

Thylacoleo is thought to have become extinct around 40,000 years ago as part of the Late Pleistocene megafauna extinctions, essentially simultanteously with the vast majority of Australian megafauna. It has been contested as to the relative importance of climatic change vs the impact of recently arrived Indigenous Australians (who arrived in Australia around 50-60,000 years ago) in the extinctions. There is limited evidence of human interaction with extinct megafauna in Australia.^[39]

References

^ Alloing-Séguier, Léanie; Sánchez-Villagra, Marcelo R.; Lee, Michael S. Y.; Lebrun, Renaud (2013). "The Bony Labyrinth in Diprotodontian Marsupial Mammals: Diversity in Extant and Extinct Forms and Relationships with Size and Phylogeny". Journal of Mammalian Evolution. 20 (3): 191–198. doi:10.1007/s10914-013-9228-3. S2CID 16385939.
^ ^a ^b "XVI. On the fossil mammals of Australia.— Part I. Description of a mutilated skull of a large marsupial carnivore (Thylacoleo carnifex, Owen), from a calcareous conglomerate stratum, eighty miles S. W. Of Melbourne, Victoria". Philosophical Transactions of the Royal Society of London. 149: 309–322. 1859. doi:10.1098/rstl.1859.0016. S2CID 110651400.
^ ^a ^b Musser, Anne (29 November 2018). "Thylacoleo carnifex". Australian Museum. Retrieved 7 May 2019.
^ ^a ^b ^c "XV. On the affinities of thylacoleo". Philosophical Transactions of the Royal Society of London. 174: 575–582. 1883. doi:10.1098/rstl.1883.0015. ISSN 0261-0523. S2CID 111347165.
^ Gill, E.D. (25 February 1973). "Antipodal distribution of the holotype bones of Thylacoleo carnifex Owen (Marsupialia)". Science Reports of the Tohoku University. Second Series, Geology. 6: 497–499. Retrieved 13 August 2020.
^ Woods, J. T. (1956). "The skull of Thylacoleo carnifex". Memoirs of the Queensland Museum. 13: 125–140.
^ Bartholomai, Alan (1962). "A new species of Thylacoleo and notes on some caudal vertebrae of Palorchestes azael". Memoirs of the Queensland Museum. 14: 33–40.
^ ^a ^b Pledge, N.S. (1977). "A new species of Thylacoleo (Marsupialia: Thylacoleonidae) with notes on the occurrences and distribution of Thylacoleonidae in South Australia". Records of the South Australian Museum. 17: 277–283.
^ Long, J.A., Archer, M., Flannery, T. & Hand, S. (2002). Prehistoric Mammals of Australia and New Guinea - 100 million Years of Evolution. Baltimore: Johns Hopkins University Press. pp. 224pp.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ Flannery, T.F.; Archer, M. (1984). "The macropodoids (Marsupialia) of the Early Pliocene Bow local fauna, central eastern New South Wales". The Australian Zoologist. 21: 357–383.
^ Pledge, Neville S. (1992). "The Curramulka local fauna: A new late Tertiary fossil assemblage from Yorke Peninsula, South Australia". The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences. 9: 115–142.
^ Naish, Darren (2004). "Of koalas and marsupial lions: the vombatiform radiation, part I". Molecular Phylogenetics and Evolution. 33 (1): 240–250. doi:10.1016/j.ympev.2004.05.004. PMID 15324852. Retrieved 17 October 2014.
^ Black, Karen; Price, Gilbert J; Archer, Michael; Hand, Suzanne J (2014). "Bearing up well? Understanding the past, present and future of Australia's koalas". Gondwana Research. 25 (3): 1186–201. Bibcode:2014GondR..25.1186B. doi:10.1016/j.gr.2013.12.008.
^ ^a ^b Black, K. H.; Archer, M.; Hand, S. J.; Godthelp, H. (2012). "The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding". In Talent, J. A. (ed.). Earth and Life. Springer Verlag. pp. 1040, 1047. doi:10.1007/978-90-481-3428-1_35. ISBN 978-90-481-3427-4.
^ Gillespie, Anna K; Archer, Michael; Hand, Suzanne J (2016). "A tiny new marsupial lion (Marsupialia, Thylacoleonidae) from the early Miocene of Australia". Palaeontologia Electronica. 19 (2). doi:10.26879/632.
^ Gillespie, Anna K.; Archer, Michael; Hand, Suzanne J. (2019-09-03). "Lekaneleo, a new genus of marsupial lion (Marsupialia, Thylacoleonidae) from the Oligocene–Miocene of Australia, and the craniodental morphology of L. roskellyae, comb. nov". Journal of Vertebrate Paleontology. 39 (5): e1703722. Bibcode:2019JVPal..39E3722G. doi:10.1080/02724634.2019.1703722. ISSN 0272-4634. S2CID 214332715.
^ ^a ^b Wroe, S; Myers, T. J; Wells, R. T; Gillespie, A (1999). "Estimating the weight of the Pleistocene marsupial lion, Thylacoleo carnifex (Thylacoleonidae:Marsupialia): Implications for the ecomorphology of a marsupial super-predator and hypotheses of impoverishment of Australian marsupial carnivore faunas". Australian Journal of Zoology. 47 (5): 489–98. doi:10.1071/ZO99006.
^ Wroe, Stephen; Myers, Troy; Seebacher, Frank; Kear, Ben; Gillespie, Anna; Crowther, Mathew; Salisbury, Steve (Summer 2003). "An alternative method for predicting body mass: the case of the Pleistocene marsupial lion". Paleobiology. 29 (3): 403–411. Bibcode:2003Pbio...29..403W. doi:10.1666/0094-8373(2003)029<0403:AAMFPB>2.0.CO;2. ISSN 0094-8373. Retrieved 14 August 2024 – via Cambridge Core.
^ ^a ^b Werdelin, L (1988). "Circumventing a Constraint - the Case of Thylacoleo (Marsupialia, Thylacoleonidae)". Australian Journal of Zoology. 36 (5): 565. doi:10.1071/ZO9880565. ISSN 0004-959X.
^ ^a ^b Wroe, S.; McHenry, C.; Thomason, J. (2005). "Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa". Proceedings of the Royal Society B: Biological Sciences. 272 (1563): 619–625. doi:10.1098/rspb.2004.2986. PMC 1564077. PMID 15817436.
^ ^a ^b Gillespie, Anna K. (2023-01-16). "Two new marsupial lion taxa (Marsupialia, Thylacoleonidae) from the early and Middle Miocene of Australia". Alcheringa: An Australasian Journal of Palaeontology. 47 (4): 506–521. Bibcode:2023Alch...47..506G. doi:10.1080/03115518.2022.2152096. ISSN 0311-5518. S2CID 256157821.
^ Wells, Roderick T.; Murray, Peter F.; Bourne, Steven J. (2009). "Pedal morphology of the marsupial lion Thylacoleo carnifex (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia". Journal of Vertebrate Paleontology. 29 (4): 1335–1340. Bibcode:2009JVPal..29.1335W. doi:10.1671/039.029.0424. S2CID 86460654.
^ "Discovery and Interpretation". Natural Worlds. Retrieved 7 May 2019.
^ Switek, Brian (31 August 2007). "Thylacoleo carnifex, ancient Australia's marsupial lion". Laelaps. Retrieved 8 May 2019.
^ ^a ^b ^c ^d Figueirido, Borja; Martín-Serra, Alberto; Janis, Christine M. (2016). "Ecomorphological determinations in the absence of living analogues: The predatory behavior of the marsupial lion (Thylacoleo carnifex) as revealed by elbow joint morphology" (PDF). Paleobiology. 42 (3): 508–531. Bibcode:2016Pbio...42..508F. doi:10.1017/pab.2015.55. hdl:1983/1f3ac566-0b08-48fb-b27c-63fe4256c138. S2CID 87168573.
^ "Extinct Marsupial Lion Tops African Lion In Fight To Death", Science Daily, 17 January 2008.
^ Dayton, Leigh (18 January 2008). "Marsupial lion was fast killer". The Australian. Archived from the original on 17 April 2009.
^ Wells, R.T.; Murray, P.F.; Bourne, S.J. (2009). "Pedal morphology of the marsupial lion Thylacoleo carnifex (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia". Journal of Vertebrate Paleontology. 29 (4): 1335–1340. Bibcode:2009JVPal..29.1335W. doi:10.1671/039.029.0424. S2CID 86460654.
^ ^a ^b Arman, Samuel D.; Prideaux, Gavin J. (15 February 2016). "Behaviour of the Pleistocene marsupial lion deduced from claw marks in a southwestern Australian cave". Scientific Reports. 6: 21372. Bibcode:2016NatSR...621372A. doi:10.1038/srep21372. PMC 4753435. PMID 26876952. S2CID 3548956.
^ ^a ^b ^c ^d Evans, A. R.; Wells, R. T.; Camens, A. B. (2018). "New skeletal material sheds light on the palaeobiology of the Pleistocene marsupial carnivore, Thylacoleo carnifex". PLOS ONE. 13 (12): e0208020. Bibcode:2018PLoSO..1308020W. doi:10.1371/journal.pone.0208020. PMC 6291118. PMID 30540785.
^ ^a ^b Switek, Brian. "Thylacoleo: Herbivore or Carnivore?". Wired.
^ ^a ^b ^c Horton, D. R.; Wright, R. V. S. (1981). "Cuts on Lancefield Bones: Carnivorous Thylacoleo, Not Humans, the Cause". Archaeology in Oceania. 16 (2): 73–80. doi:10.1002/j.1834-4453.1981.tb00009.x. JSTOR 40386545.
^ De Vis, C. W. (1883). "On tooth-marked bones of extinct marsupials". Proceedings of the Linnean Society of New South Wales. 8: 187–190. doi:10.5962/bhl.part.28646.
^ Anderson, C. (1929). "Palaeontological notes no. 1. Macropus titan Owen and Thylacoleo carnifex Owen". Records of the Australian Museum. 17: 35–49. doi:10.3853/j.0067-1975.17.1929.752.
^ Tyndale-Biscoe, Hugh (2005). Life of marsupials. Collingwood, Vic.: CSIRO. ISBN 978-0-643-09220-4.^{[page needed]}
^ "Marsupial lion 'could climb trees'". BBC News. 2016-02-15.
^ Prideaux, Gavin J.; Long, John A.; Ayliffe, Linda K.; Hellstrom, John C.; Pillans, Brad; Boles, Walter E.; Hutchinson, Mark N.; Roberts, Richard G.; Cupper, Matthew L.; Arnold, Lee J.; Devine, Paul D.; Warburton, Natalie M. (January 2007). "An arid-adapted middle Pleistocene vertebrate fauna from south-central Australia". Nature. 445 (7126): 422–425. Bibcode:2007Natur.445..422P. doi:10.1038/nature05471. ISSN 0028-0836. PMID 17251978.
^ Case, J. A. (1985-01-01). "Differences in prey utilization by Pleistocene marsupial carnivores, Thylacoleo carnifex (Thylacoleonidae) and Thylacinus cynocephalus (Thylacinidae)". Australian Mammalogy. 8 (1): 45–52. doi:10.1071/AM85002. ISSN 0310-0049.
^ Hocknull, Scott A.; Lewis, Richard; Arnold, Lee J.; Pietsch, Tim; Joannes-Boyau, Renaud; Price, Gilbert J.; Moss, Patrick; Wood, Rachel; Dosseto, Anthony; Louys, Julien; Olley, Jon; Lawrence, Rochelle A. (2020-05-18). "Extinction of eastern Sahul megafauna coincides with sustained environmental deterioration". Nature Communications. 11 (1): 2250. Bibcode:2020NatCo..11.2250H. doi:10.1038/s41467-020-15785-w. ISSN 2041-1723. PMC 7231803. PMID 32418985.

External links

[1] Alloing-Séguier, Léanie; Sánchez-Villagra, Marcelo R.; Lee, Michael S. Y.; Lebrun, Renaud (2013). "The Bony Labyrinth in Diprotodontian Marsupial Mammals: Diversity in Extant and Extinct Forms and Relationships with Size and Phylogeny". Journal of Mammalian Evolution. 20 (3): 191–198. doi:10.1007/s10914-013-9228-3. S2CID 16385939.

[Owen1859XVI-2] "XVI. On the fossil mammals of Australia.— Part I. Description of a mutilated skull of a large marsupial carnivore (Thylacoleo carnifex, Owen), from a calcareous conglomerate stratum, eighty miles S. W. Of Melbourne, Victoria". Philosophical Transactions of the Royal Society of London. 149: 309–322. 1859. doi:10.1098/rstl.1859.0016. S2CID 110651400.

[:4-3] Musser, Anne (29 November 2018). "Thylacoleo carnifex". Australian Museum. Retrieved 7 May 2019.

[Owen1883XV-4] "XV. On the affinities of thylacoleo". Philosophical Transactions of the Royal Society of London. 174: 575–582. 1883. doi:10.1098/rstl.1883.0015. ISSN 0261-0523. S2CID 111347165.

[5] Gill, E.D. (25 February 1973). "Antipodal distribution of the holotype bones of Thylacoleo carnifex Owen (Marsupialia)". Science Reports of the Tohoku University. Second Series, Geology. 6: 497–499. Retrieved 13 August 2020.

[Woods1956-6] Woods, J. T. (1956). "The skull of Thylacoleo carnifex". Memoirs of the Queensland Museum. 13: 125–140.

[Bartholomai1962-7] Bartholomai, Alan (1962). "A new species of Thylacoleo and notes on some caudal vertebrae of Palorchestes azael". Memoirs of the Queensland Museum. 14: 33–40.

[Pledge1977-8] Pledge, N.S. (1977). "A new species of Thylacoleo (Marsupialia: Thylacoleonidae) with notes on the occurrences and distribution of Thylacoleonidae in South Australia". Records of the South Australian Museum. 17: 277–283.

[9] Long, J.A., Archer, M., Flannery, T. & Hand, S. (2002). Prehistoric Mammals of Australia and New Guinea - 100 million Years of Evolution. Baltimore: Johns Hopkins University Press. pp. 224pp.{{cite book}}: CS1 maint: multiple names: authors list (link)

[Flannery1984-10] Flannery, T.F.; Archer, M. (1984). "The macropodoids (Marsupialia) of the Early Pliocene Bow local fauna, central eastern New South Wales". The Australian Zoologist. 21: 357–383.

[Pledge1992-11] Pledge, Neville S. (1992). "The Curramulka local fauna: A new late Tertiary fossil assemblage from Yorke Peninsula, South Australia". The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences. 9: 115–142.

[Scientific_American-12] Naish, Darren (2004). "Of koalas and marsupial lions: the vombatiform radiation, part I". Molecular Phylogenetics and Evolution. 33 (1): 240–250. doi:10.1016/j.ympev.2004.05.004. PMID 15324852. Retrieved 17 October 2014.

[Black_et_al_2014-13] Black, Karen; Price, Gilbert J; Archer, Michael; Hand, Suzanne J (2014). "Bearing up well? Understanding the past, present and future of Australia's koalas". Gondwana Research. 25 (3): 1186–201. Bibcode:2014GondR..25.1186B. doi:10.1016/j.gr.2013.12.008.

[Black2012-14] Black, K. H.; Archer, M.; Hand, S. J.; Godthelp, H. (2012). "The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding". In Talent, J. A. (ed.). Earth and Life. Springer Verlag. pp. 1040, 1047. doi:10.1007/978-90-481-3428-1_35. ISBN 978-90-481-3427-4.

[15] Gillespie, Anna K; Archer, Michael; Hand, Suzanne J (2016). "A tiny new marsupial lion (Marsupialia, Thylacoleonidae) from the early Miocene of Australia". Palaeontologia Electronica. 19 (2). doi:10.26879/632.

[16] Gillespie, Anna K.; Archer, Michael; Hand, Suzanne J. (2019-09-03). "Lekaneleo, a new genus of marsupial lion (Marsupialia, Thylacoleonidae) from the Oligocene–Miocene of Australia, and the craniodental morphology of L. roskellyae, comb. nov". Journal of Vertebrate Paleontology. 39 (5): e1703722. Bibcode:2019JVPal..39E3722G. doi:10.1080/02724634.2019.1703722. ISSN 0272-4634. S2CID 214332715.

[:2-17] Wroe, S; Myers, T. J; Wells, R. T; Gillespie, A (1999). "Estimating the weight of the Pleistocene marsupial lion, Thylacoleo carnifex (Thylacoleonidae:Marsupialia): Implications for the ecomorphology of a marsupial super-predator and hypotheses of impoverishment of Australian marsupial carnivore faunas". Australian Journal of Zoology. 47 (5): 489–98. doi:10.1071/ZO99006.

[18] Wroe, Stephen; Myers, Troy; Seebacher, Frank; Kear, Ben; Gillespie, Anna; Crowther, Mathew; Salisbury, Steve (Summer 2003). "An alternative method for predicting body mass: the case of the Pleistocene marsupial lion". Paleobiology. 29 (3): 403–411. Bibcode:2003Pbio...29..403W. doi:10.1666/0094-8373(2003)029<0403:AAMFPB>2.0.CO;2. ISSN 0094-8373. Retrieved 14 August 2024 – via Cambridge Core.

[:0-19] Werdelin, L (1988). "Circumventing a Constraint - the Case of Thylacoleo (Marsupialia, Thylacoleonidae)". Australian Journal of Zoology. 36 (5): 565. doi:10.1071/ZO9880565. ISSN 0004-959X.

[rspb.royalsocietypublishing.org-20] Wroe, S.; McHenry, C.; Thomason, J. (2005). "Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa". Proceedings of the Royal Society B: Biological Sciences. 272 (1563): 619–625. doi:10.1098/rspb.2004.2986. PMC 1564077. PMID 15817436.

[:3-21] Gillespie, Anna K. (2023-01-16). "Two new marsupial lion taxa (Marsupialia, Thylacoleonidae) from the early and Middle Miocene of Australia". Alcheringa: An Australasian Journal of Palaeontology. 47 (4): 506–521. Bibcode:2023Alch...47..506G. doi:10.1080/03115518.2022.2152096. ISSN 0311-5518. S2CID 256157821.

[22] Wells, Roderick T.; Murray, Peter F.; Bourne, Steven J. (2009). "Pedal morphology of the marsupial lion Thylacoleo carnifex (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia". Journal of Vertebrate Paleontology. 29 (4): 1335–1340. Bibcode:2009JVPal..29.1335W. doi:10.1671/039.029.0424. S2CID 86460654.

[:1-23] "Discovery and Interpretation". Natural Worlds. Retrieved 7 May 2019.

[24] Switek, Brian (31 August 2007). "Thylacoleo carnifex, ancient Australia's marsupial lion". Laelaps. Retrieved 8 May 2019.

[figu2016-25] Figueirido, Borja; Martín-Serra, Alberto; Janis, Christine M. (2016). "Ecomorphological determinations in the absence of living analogues: The predatory behavior of the marsupial lion (Thylacoleo carnifex) as revealed by elbow joint morphology" (PDF). Paleobiology. 42 (3): 508–531. Bibcode:2016Pbio...42..508F. doi:10.1017/pab.2015.55. hdl:1983/1f3ac566-0b08-48fb-b27c-63fe4256c138. S2CID 87168573.

[26] "Extinct Marsupial Lion Tops African Lion In Fight To Death", Science Daily, 17 January 2008.

[27] Dayton, Leigh (18 January 2008). "Marsupial lion was fast killer". The Australian. Archived from the original on 17 April 2009.

[28] Wells, R.T.; Murray, P.F.; Bourne, S.J. (2009). "Pedal morphology of the marsupial lion Thylacoleo carnifex (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia". Journal of Vertebrate Paleontology. 29 (4): 1335–1340. Bibcode:2009JVPal..29.1335W. doi:10.1671/039.029.0424. S2CID 86460654.

[Claw-29] Arman, Samuel D.; Prideaux, Gavin J. (15 February 2016). "Behaviour of the Pleistocene marsupial lion deduced from claw marks in a southwestern Australian cave". Scientific Reports. 6: 21372. Bibcode:2016NatSR...621372A. doi:10.1038/srep21372. PMC 4753435. PMID 26876952. S2CID 3548956.

[Evans2018-30] Evans, A. R.; Wells, R. T.; Camens, A. B. (2018). "New skeletal material sheds light on the palaeobiology of the Pleistocene marsupial carnivore, Thylacoleo carnifex". PLOS ONE. 13 (12): e0208020. Bibcode:2018PLoSO..1308020W. doi:10.1371/journal.pone.0208020. PMC 6291118. PMID 30540785.

[wired2009-31] Switek, Brian. "Thylacoleo: Herbivore or Carnivore?". Wired.

[bite-32] Horton, D. R.; Wright, R. V. S. (1981). "Cuts on Lancefield Bones: Carnivorous Thylacoleo, Not Humans, the Cause". Archaeology in Oceania. 16 (2): 73–80. doi:10.1002/j.1834-4453.1981.tb00009.x. JSTOR 40386545.

[scavenger-33] De Vis, C. W. (1883). "On tooth-marked bones of extinct marsupials". Proceedings of the Linnean Society of New South Wales. 8: 187–190. doi:10.5962/bhl.part.28646.

[melons-34] Anderson, C. (1929). "Palaeontological notes no. 1. Macropus titan Owen and Thylacoleo carnifex Owen". Records of the Australian Museum. 17: 35–49. doi:10.3853/j.0067-1975.17.1929.752.

[35] Tyndale-Biscoe, Hugh (2005). Life of marsupials. Collingwood, Vic.: CSIRO. ISBN 978-0-643-09220-4.^{[page needed]}

[36] "Marsupial lion 'could climb trees'". BBC News. 2016-02-15.

[37] Prideaux, Gavin J.; Long, John A.; Ayliffe, Linda K.; Hellstrom, John C.; Pillans, Brad; Boles, Walter E.; Hutchinson, Mark N.; Roberts, Richard G.; Cupper, Matthew L.; Arnold, Lee J.; Devine, Paul D.; Warburton, Natalie M. (January 2007). "An arid-adapted middle Pleistocene vertebrate fauna from south-central Australia". Nature. 445 (7126): 422–425. Bibcode:2007Natur.445..422P. doi:10.1038/nature05471. ISSN 0028-0836. PMID 17251978.

[38] Case, J. A. (1985-01-01). "Differences in prey utilization by Pleistocene marsupial carnivores, Thylacoleo carnifex (Thylacoleonidae) and Thylacinus cynocephalus (Thylacinidae)". Australian Mammalogy. 8 (1): 45–52. doi:10.1071/AM85002. ISSN 0310-0049.

[39] Hocknull, Scott A.; Lewis, Richard; Arnold, Lee J.; Pietsch, Tim; Joannes-Boyau, Renaud; Price, Gilbert J.; Moss, Patrick; Wood, Rachel; Dosseto, Anthony; Louys, Julien; Olley, Jon; Lawrence, Rochelle A. (2020-05-18). "Extinction of eastern Sahul megafauna coincides with sustained environmental deterioration". Nature Communications. 11 (1): 2250. Bibcode:2020NatCo..11.2250H. doi:10.1038/s41467-020-15785-w. ISSN 2041-1723. PMC 7231803. PMID 32418985.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

@@ Line 13: / Line 13: @@
 }}
-'''''Thylacoleo''''' ("pouch lion") is an [[Extinction|extinct]] [[genus]] of carnivorous [[marsupial]]s that lived in [[Australia]] from the late [[Pliocene]] to the [[Late Pleistocene]] (until around 40,000 years ago), often known as '''marsupial lions'''. They were the largest and last members of the family [[Thylacoleonidae]], occupying the position of [[apex predator]] within Australian ecosystems. The largest and last species, ''Thylacoleo carnifex'' approached the weight of a [[lion]]ess. The estimated average weight for the species ranges from {{convert|101|to|130|kg|abbr=on}}.<ref>{{cite journal | doi=10.1007/s10914-013-9228-3 | volume=20 | issue=3 | title=The Bony Labyrinth in Diprotodontian Marsupial Mammals: Diversity in Extant and Extinct Forms and Relationships with Size and Phylogeny | journal=Journal of Mammalian Evolution | pages=191–198|year = 2013|last1 = Alloing-Séguier|first1 = Léanie| last2=Sánchez-Villagra | first2=Marcelo R. | last3=Lee | first3=Michael S. Y. | last4=Lebrun | first4=Renaud | s2cid=16385939 }}</ref>
+'''''Thylacoleo''''' ("pouch lion") is an [[Extinction|extinct]] [[genus]] of carnivorous [[marsupial]]s that lived in [[Australia]] from the late [[Pliocene]] to the [[Late Pleistocene]] (until around 40,000 years ago), often known as '''marsupial lions'''. They were the largest and last members of the family [[Thylacoleonidae]], occupying the position of [[apex predator]] within Australian ecosystems. The largest and last species, ''Thylacoleo carnifex'', approached the weight of a [[lion]]ess. The estimated average weight for the species ranges from {{convert|101|to|130|kg|abbr=on}}.<ref>{{cite journal | doi=10.1007/s10914-013-9228-3 | volume=20 | issue=3 | title=The Bony Labyrinth in Diprotodontian Marsupial Mammals: Diversity in Extant and Extinct Forms and Relationships with Size and Phylogeny | journal=Journal of Mammalian Evolution | pages=191–198|year = 2013|last1 = Alloing-Séguier|first1 = Léanie| last2=Sánchez-Villagra | first2=Marcelo R. | last3=Lee | first3=Michael S. Y. | last4=Lebrun | first4=Renaud | s2cid=16385939 }}</ref>
 ==Taxonomy==
+The first ''Thylacoleo'' fossil findings, discovered by [[Thomas Mitchell (explorer)|Thomas Mitchell]] were found in the 1830s in the Wellington Valley of New South Wales, though not recognised as such at the time. The generic holotype, consisting of broken teeth, jaws, and a skull was discovered by a pastoralist, William Avery, near Lake Colungolac from which the species ''Thylacoleo carnifex'' was described by [[Richard Owen]].<ref name="Owen1859XVI">{{cite journal |year=1859 |title=XVI. On the fossil mammals of Australia.— Part I. Description of a mutilated skull of a large marsupial carnivore (Thylacoleo carnifex, Owen), from a calcareous conglomerate stratum, eighty miles S. W. Of Melbourne, Victoria |journal=Philosophical Transactions of the Royal Society of London |volume=149 |pages=309–322 |doi=10.1098/rstl.1859.0016 |s2cid=110651400}}</ref> It was not until 1966 that the first nearly-complete skeleton was found. The only pieces missing were a foot and the tail. Currently, the [[Nullarbor Plain]] of West Australia remains to be the greatest finding site. These fossils now reside at the Australian Museum.<ref name=":4">{{Cite web |last=Musser |first=Anne |date=29 November 2018 |title=Thylacoleo carnifex |url=http://australianmuseum.net.au/Thylacoleo-carnifex |access-date=7 May 2019 |website=Australian Museum}}</ref><ref name="Owen1883XV" />
 The genus was first published in 1859, erected to describe the type species ''Thylacoleo carnifex''. The new taxon was established in examination of fossil specimens provided to [[Richard Owen]]. The familial alliance takes its name from this description, the so-called marsupial lions of [[Thylacoleonidae]].
-The colloquial name "marsupial lion" alludes to the superficial resemblance to the placental lion and its ecological niche as a large predator. Contrary to its common name, ''Thylacoleo'' is not closely related to the modern lion (''[[Lion|Panthera leo]]'').
+The colloquial name "marsupial lion" alludes to the genus name, which was named after its superficial resemblance to the placental lion and its ecological niche as a large predator.
-'''Genus:''' ''Thylacoleo'' (''Thylacopardus'') – Australia's marsupial lions, that lived from about 2 million years ago, during the Late Pliocene Epoch and became extinct about 30,000 years ago{{Citation needed|date=April 2021}}, during the Late Pleistocene Epoch. Three species are known:
+'''Genus:''' ''Thylacoleo'' (''Thylacopardus'') – Australia's marsupial lions, that lived from about 2&nbsp;million years ago, during the Late Pliocene Epoch and became extinct about 40,000 years ago, during the [[Late Pleistocene]] Epoch. Three species are known:
-*''Thylacoleo carnifex'' The holotype cranium was collected from [[Western District Lakes|Lake Colongulac]] in 1843 by pastoralist William Adeney. A partial rostrum collected by Adeney in 1876 from the same locality would later be found to belong to the same individual.<ref>{{cite journal |last1= Gill |first1= E.D. |date=25 February 1973 |title= Antipodal distribution of the holotype bones of ''Thylacoleo carnifex'' Owen (Marsupialia) |journal= Science Reports of the Tohoku University. Second Series, Geology |volume= 6 |pages= 497–499 |url = https://tohoku.repo.nii.ac.jp/?action=repository_uri&item_id=10862&file_id=18&file_no=1 |access-date= 13 August 2020}}</ref>
+*''Thylacoleo carnifex'' The holotype cranium was collected from [[Western District Lakes|Lake Colongulac]] in 1843 by pastoralist William Adeney. A partial rostrum collected by Adeney in 1876 from the same locality would later be found to belong to the same individual.<ref>{{cite journal |last1= Gill |first1= E.D. |date=25 February 1973 |title= Antipodal distribution of the holotype bones of ''Thylacoleo carnifex'' Owen (Marsupialia) |journal= Science Reports of the Tohoku University. Second Series, Geology |volume= 6 |pages= 497–499 |url = https://tohoku.repo.nii.ac.jp/?action=repository_uri&item_id=10862&file_id=18&file_no=1 |access-date= 13 August 2020}}</ref> It was not until 1966 that the first nearly-complete skeleton was found.<ref name=":4" /><ref name="Owen1883XV" />
-*''Thylacoleo crassidentatus'' lived during the [[Pliocene]], around 5 million years ago, and was about the size of a large dog. Its fossils have been found in southeastern [[Queensland]].<ref name="Woods1956">{{cite journal |last1=Woods |first1=J. T. |title=The skull of Thylacoleo carnifex |journal=Memoirs of the Queensland Museum |date=1956 |volume=13 |pages=125–140 |url=https://www.biodiversitylibrary.org/part/303455}}</ref><ref name="Bartholomai1962">{{cite journal |last1=Bartholomai |first1=Alan |title=A new species of Thylacoleo and notes on some caudal vertebrae of Palorchestes azael |journal=Memoirs of the Queensland Museum |date=1962 |volume=14 |pages=33–40 |url=https://www.biodiversitylibrary.org/part/303461}}</ref>
+*''Thylacoleo crassidentatus'' lived during the [[Pliocene]], around 5&nbsp;million years ago, and was about the size of a large dog. Its fossils have been found in southeastern [[Queensland]].<ref name="Woods1956">{{cite journal |last1=Woods |first1=J. T. |title=The skull of Thylacoleo carnifex |journal=Memoirs of the Queensland Museum |date=1956 |volume=13 |pages=125–140 |url=https://www.biodiversitylibrary.org/part/303455}}</ref><ref name="Bartholomai1962">{{cite journal |last1=Bartholomai |first1=Alan |title=A new species of Thylacoleo and notes on some caudal vertebrae of Palorchestes azael |journal=Memoirs of the Queensland Museum |date=1962 |volume=14 |pages=33–40 |url=https://www.biodiversitylibrary.org/part/303461}}</ref>
-*''Thylacoleo hilli'' lived during the Pliocene and was half the size of ''T. crassidentatus''. It is the oldest member of the genus.<ref name="Pledge1977"/>
+*''Thylacoleo hilli'' lived during the Pliocene and was half the size of ''T.&nbsp;crassidentatus''. It is the oldest member of the genus.<ref name="Pledge1977"/>
-Fossils of other representatives of Thylacoleonidae, such as ''[[Microleo]]'' and ''[[Wakaleo]]'', date back to the Late Oligocene Epoch, some 24 million years ago.<ref>{{cite book | author = Long, J.A., Archer, M., Flannery, T. & Hand, S. | year = 2002 | title = Prehistoric Mammals of Australia and New Guinea - 100 million Years of Evolution | publisher = Johns Hopkins University Press | location = Baltimore | pages = 224pp}}</ref>
+Fossils of other representatives of Thylacoleonidae, such as ''[[Microleo]]'' and ''[[Wakaleo]]'', date back to the Late Oligocene Epoch, some 24&nbsp;million years ago.<ref>{{cite book | author = Long, J.A., Archer, M., Flannery, T. & Hand, S. | year = 2002 | title = Prehistoric Mammals of Australia and New Guinea - 100 million Years of Evolution | publisher = Johns Hopkins University Press | location = Baltimore | pages = 224pp}}</ref>
-''T. hilli'' was described by [[Neville Pledge]] in a study published in the records of the South Australia Museum in 1977. The holotype is a third premolar, discovered at a cave in [[Curramulka]] in South Australia, exhibiting the carnivorous characteristics of the genus and around half the size of ''T. carnifex''. This tooth was collected by Alan Hill, a speleologist and founding member of the Cave Exploration Group of South Australia, while examining a site known as the "Town Cave" in 1956; the specific epithet ''hilli'' honours the collector of the first specimen.<ref name="Pledge1977">{{cite journal |last1=Pledge |first1=N.S. |authorlink1=Neville S. Pledge |title=A new species of ''Thylacoleo'' (Marsupialia: Thylacoleonidae) with notes on the occurrences and distribution of Thylacoleonidae in South Australia |journal=Records of the South Australian Museum |date=1977 |volume=17 |pages=277–283 |url=https://www.biodiversitylibrary.org/part/78709}}</ref> Material found amidst the fauna at [[Bow River (New South Wales)|Bow River]] in New South Wales, dated to the early [[Pliocene]], was also referred to the species in 1982.
+''T. hilli'' was described by [[Neville Pledge]] in a study published in the records of the South Australia Museum in 1977. The holotype is a third premolar, discovered at a cave in [[Curramulka]] in South Australia, exhibiting the carnivorous characteristics of the genus and around half the size of ''T. carnifex''. This tooth was collected by Alan Hill, a speleologist and founding member of the Cave Exploration Group of South Australia, while examining a site known as the "Town Cave" in 1956; the specific epithet ''hilli'' honours the collector of the first specimen.<ref name="Pledge1977">{{cite journal |last1=Pledge |first1=N.S. |authorlink1=Neville S. Pledge |title=A new species of ''Thylacoleo'' (Marsupialia: Thylacoleonidae) with notes on the occurrences and distribution of Thylacoleonidae in South Australia |journal=Records of the South Australian Museum |date=1977 |volume=17 |pages=277–283 |url=https://www.biodiversitylibrary.org/part/78709}}</ref> Material found amidst the fauna at [[Bow River (New South Wales)|Bow River]] in New South Wales, dated to the early [[Pliocene]], was also referred to the species in 1982.<ref name="Flannery1984">{{cite journal |last1=Flannery |first1=T.F. |last2=Archer |first2=M. |title=The macropodoids (Marsupialia) of the Early Pliocene Bow local fauna, central eastern New South Wales |journal=The Australian Zoologist |date=1984 |volume=21 |pages=357–383 |url=https://www.biodiversitylibrary.org/part/50064}}</ref>
-<ref name="Flannery1984">{{cite journal |last1=Flannery |first1=T.F. |last2=Archer |first2=M. |title=The macropodoids (Marsupialia) of the Early Pliocene Bow local fauna, central eastern New South Wales |journal=The Australian Zoologist |date=1984 |volume=21 |pages=357–383 |url=https://www.biodiversitylibrary.org/part/50064}}</ref>
+A fragment of an incisor, unworn and only diagnosable to the genus, was located at a site in Curramulka, close to the Town Cave site, and referred to the species for the apparent correlation in size when compared to the better known ''T.&nbsp;carnifex''.<ref name="Pledge1992">{{cite journal |last1=Pledge |first1=Neville S. |title=The Curramulka local fauna: A new late Tertiary fossil assemblage from Yorke Peninsula, South Australia |journal=The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences |date=1992 |volume=9 |pages=115–142 |url=https://www.biodiversitylibrary.org/part/263122}}</ref>
-A fragment of an incisor, unworn and only diagnosable to the genus, was located at a site in Curramulka, close to the Town Cave site, and referred to the species for the apparent correlation in size when compared to the better known ''T. carnifex''.<ref name="Pledge1992">{{cite journal |last1=Pledge |first1=Neville S. |title=The Curramulka local fauna: A new late Tertiary fossil assemblage from Yorke Peninsula, South Australia |journal=The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences |date=1992 |volume=9 |pages=115–142 |url=https://www.biodiversitylibrary.org/part/263122}}</ref>
 The marsupial lion is classified in the order Diprotodontia along with many other well-known marsupials such as kangaroos, [[Phalangeriformes|possums]], and the [[koala]]. It is further classified in its own [[family (biology)|family]], the [[Thylacoleonidae]], of which three [[genus|genera]] and 11 [[species]] are recognised, all extinct. The term marsupial lion (lower case) is often applied to other members of this family. Distinct possum-like characteristics led ''Thylacoleo'' to be regarded as members of [[Phalangeroidea]] for a few decades. Though a few authors continued to hint at phalangeroid affinities for thylacoleonids as recently as the 1990s, cranial and other characters have generally led to their inclusion within [[vombatiformes]], and as stem-members of the wombat lineage.<ref name="Scientific American">{{cite journal |last1=Naish |first1=Darren |year=2004 |title=Of koalas and marsupial lions: the vombatiform radiation, part I |url=http://blogs.scientificamerican.com/tetrapod-zoology/2011/10/26/vombatiform-radiation-part-i/ |journal=Molecular Phylogenetics and Evolution |volume=33 |issue=1 |pages=240–250 |doi=10.1016/j.ympev.2004.05.004 |pmid=15324852 |accessdate=17 October 2014}}</ref> Marsupial lions and other ecologically and morphologically diverse vombatiforms were once represented by over 60 species of [[carnivorous]], [[herbivorous]], [[Terrestrial animal|terrestrial]] and [[arboreal]] forms ranging in size from 3&nbsp;kg to 2.5 tonnes. Only two families represented by four [[herbivorous]] species (koalas and three species of wombat) have survived into modern times and are considered the marsupial lion's closest living relatives.<ref name="Black et al 2014">{{cite journal |doi=10.1016/j.gr.2013.12.008 |title=Bearing up well? Understanding the past, present and future of Australia's koalas |journal=Gondwana Research |volume=25 |issue=3 |pages=1186–201 |year=2014 |last1=Black |first1=Karen |authorlink1 = Karen H. Black |last2=Price |first2=Gilbert J |last3=Archer |first3=Michael |last4=Hand |first4=Suzanne J |bibcode=2014GondR..25.1186B }}</ref>
 ==Evolution==
-The ancestors of [[thylacoleonid]]s are believed to have been [[herbivore]]s, something unusual for carnivores. They are members of the [[Vombatiformes]], an almost entirely herbivorous order of marsupials, the only extant representatives of which are [[koala]]s and [[wombat]]s, as well as extinct members such as the [[diprotodontids]] and [[palorchestids]].<ref name="Black2012">{{cite book |last1=Black |first1=K. H. |url=https://www.researchgate.net/publication/259220987 |last2=Archer |first2=M. |last3=Hand |first3=S. J. |last4=Godthelp |first4=H. |title=Earth and Life |publisher=Springer Verlag |year=2012 |editor-last=Talent |editor-first=J. A. |pages=1040, 1047 |chapter=The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding |doi=10.1007/978-90-481-3428-1_35 |isbn=978-90-481-3427-4 |authorlink=Karen H. Black}}</ref> The group first appeared in the Late [[Oligocene]]. The earliest thylacoleonids like ''[[Microleo]]'' were small possum-like animals,<ref>{{cite journal |doi=10.26879/632 |title=A tiny new marsupial lion (Marsupialia, Thylacoleonidae) from the early Miocene of Australia |journal=Palaeontologia Electronica |volume=19 |issue=2 |year=2016 |last1=Gillespie |first1=Anna K |last2=Archer |first2=Michael |last3=Hand |first3=Suzanne J |doi-access=free }}</ref> with the group increasing in size during the [[Miocene]], with representatives like the leopoard sized ''[[Wakaleo]].'' The genus ''Thylacoleo'' first appeared during the [[Pliocene]], and represented the only extant genus of the family from that time until the end of the [[Pleistocene]]. The youngest representative of ''Thylacoleo'' and the thylacoleonids, ''T. carnifex'', is the largest known member of the family.<ref name="Black2012" /> The earliest thylacoleonids are thought to have been arboreal (tree dwelling) animals,<ref>{{Cite journal |last=Gillespie |first=Anna K. |last2=Archer |first2=Michael |last3=Hand |first3=Suzanne J. |date=2019-09-03 |title=Lekaneleo , a new genus of marsupial lion (Marsupialia, Thylacoleonidae) from the Oligocene–Miocene of Australia, and the craniodental morphology of L. roskellyae , comb. nov. |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2019.1703722 |journal=Journal of Vertebrate Paleontology |language=en |volume=39 |issue=5 |pages=e1703722 |doi=10.1080/02724634.2019.1703722 |issn=0272-4634}}</ref> while ''Thylacoleo'' is thought to be terrestrial with some climbing capabilities.<ref name=":2" />
+The ancestors of [[thylacoleonid]]s are believed to have been [[herbivore]]s, something unusual for carnivores. They are members of the [[Vombatiformes]], an almost entirely herbivorous order of marsupials, the only extant representatives of which are [[koala]]s and [[wombat]]s, as well as extinct members such as the [[diprotodontids]] and [[palorchestids]].<ref name="Black2012">{{cite book |last1=Black |first1=K. H. |url=https://www.researchgate.net/publication/259220987 |last2=Archer |first2=M. |last3=Hand |first3=S. J. |last4=Godthelp |first4=H. |title=Earth and Life |publisher=Springer Verlag |year=2012 |editor-last=Talent |editor-first=J. A. |pages=1040, 1047 |chapter=The Rise of Australian Marsupials: A Synopsis of Biostratigraphic, Phylogenetic, Palaeoecologic and Palaeobiogeographic Understanding |doi=10.1007/978-90-481-3428-1_35 |isbn=978-90-481-3427-4 |authorlink=Karen H. Black}}</ref> The group first appeared in the Late [[Oligocene]]. The earliest thylacoleonids like ''[[Microleo]]'' were small possum-like animals,<ref>{{cite journal |doi=10.26879/632 |title=A tiny new marsupial lion (Marsupialia, Thylacoleonidae) from the early Miocene of Australia |journal=Palaeontologia Electronica |volume=19 |issue=2 |year=2016 |last1=Gillespie |first1=Anna K |last2=Archer |first2=Michael |last3=Hand |first3=Suzanne J |doi-access=free }}</ref> with the group increasing in size during the [[Miocene]], with representatives like the leopard-sized ''[[Wakaleo]].'' The genus ''Thylacoleo'' first appeared during the [[Pliocene]], and represented the only extant genus of the family from that time until the end of the [[Pleistocene]]. The youngest representative of ''Thylacoleo'' and the thylacoleonids, ''T.&nbsp;carnifex'', is the largest known member of the family.<ref name="Black2012" /> The earliest thylacoleonids are thought to have been arboreal (tree dwelling) animals,<ref>{{Cite journal |last1=Gillespie |first1=Anna K. |last2=Archer |first2=Michael |last3=Hand |first3=Suzanne J. |date=2019-09-03 |title=Lekaneleo, a new genus of marsupial lion (Marsupialia, Thylacoleonidae) from the Oligocene–Miocene of Australia, and the craniodental morphology of L. roskellyae, comb. nov. |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2019.1703722 |journal=Journal of Vertebrate Paleontology |language=en |volume=39 |issue=5 |pages=e1703722 |doi=10.1080/02724634.2019.1703722 |bibcode=2019JVPal..39E3722G |s2cid=214332715 |issn=0272-4634}}</ref> while ''Thylacoleo'' is thought to be terrestrial with some climbing capabilities.<ref name=":2">{{cite journal |last1=Wroe |first1=S |last2=Myers |first2=T. J |last3=Wells |first3=R. T |last4=Gillespie |first4=A |year=1999 |title=Estimating the weight of the Pleistocene marsupial lion, Thylacoleo carnifex (Thylacoleonidae:Marsupialia): Implications for the ecomorphology of a marsupial super-predator and hypotheses of impoverishment of Australian marsupial carnivore faunas |journal=Australian Journal of Zoology |volume=47 |issue=5 |pages=489–98 |doi=10.1071/ZO99006}}</ref>
 ==Description==
-''T. carnifex'' is the [[Largest mammals#Marsupials .28Marsupialia.29|largest]] carnivorous [[mammal]] known to have ever existed in Australia, and one of the larger [[metatheria]]n carnivores of the world (comparable to ''[[Thylacosmilus]]'' and ''[[Borhyaena]]'' species, but smaller than [[Proborhyaenidae|''Proborhyaena gigantea'']]). Individuals ranged up to around {{convert|75|cm|in|abbr=on}} high at the shoulder and about {{convert|150|cm|in|abbr=on}} from head to tail. Measurements taken from a number of specimens show they averaged {{convert|101|to|130|kg|lb|abbr=on}} in weight, although individuals as large as {{convert|124|-|160|kg|lb|abbr=on}} might not have been uncommon, and the largest weight was of {{convert|128|-|164|kg|lb|abbr=on}}.<ref name=":2">{{cite journal |doi=10.1071/ZO99006 |title=Estimating the weight of the Pleistocene marsupial lion, Thylacoleo carnifex (Thylacoleonidae:Marsupialia): Implications for the ecomorphology of a marsupial super-predator and hypotheses of impoverishment of Australian marsupial carnivore faunas |journal=Australian Journal of Zoology |volume=47 |issue=5 |pages=489–98 |year=1999 |last1=Wroe |first1=S |last2=Myers |first2=T. J |last3=Wells |first3=R. T |last4=Gillespie |first4=A }}</ref> This would make it comparable to female lions and female [[tiger]]s in general size.
+''T. carnifex'' is the [[Largest mammals#Marsupials .28Marsupialia.29|largest]] carnivorous [[mammal]] known to have ever existed in Australia, and one of the largest [[metatheria]]n carnivores known (comparable to ''[[Thylacosmilus]]'' and ''[[Borhyaena]]'' species, but smaller than [[Proborhyaenidae|''Proborhyaena gigantea'']]). Individuals ranged up to around {{convert|75|cm|in|abbr=on}} high at the shoulder and about {{convert|150|cm|in|abbr=on}} from head to tail. Measurements taken from a number of specimens show they averaged {{convert|101|to|130|kg|lb|abbr=on}} in weight, although individuals as large as {{convert|124|-|160|kg|lb|abbr=on}} might not have been uncommon, and the largest weight was of {{convert|128|-|164|kg|lb|abbr=on}}. This would make it comparable to female lions and female [[tiger]]s in general size.<ref name=":2" /> Estimates of the size of ''T. carnifex'' based on dental remains are typically dubious, in contrast to estimates based on proximal limb bone circumference.<ref>{{Cite journal |last1=Wroe |first1=Stephen |last2=Myers |first2=Troy |last3=Seebacher |first3=Frank |last4=Kear |first4=Ben |last5=Gillespie |first5=Anna |last6=Crowther |first6=Mathew |last7=Salisbury |first7=Steve |date=Summer 2003 |title=An alternative method for predicting body mass: the case of the Pleistocene marsupial lion |url=https://www.cambridge.org/core/journals/paleobiology/article/abs/an-alternative-method-for-predicting-body-mass-the-case-of-the-pleistocene-marsupial-lion/A3DD985802D1BF1A361AFC1B7155C55B |journal=[[Paleobiology (journal)|Paleobiology]] |language=en |volume=29 |issue=3 |pages=403–411 |doi=10.1666/0094-8373(2003)029<0403:AAMFPB>2.0.CO;2 |bibcode=2003Pbio...29..403W |issn=0094-8373 |access-date=14 August 2024 |via=Cambridge Core}}</ref>
 === Skull ===
-[[Image:Megafauna.jpg|thumb|Skull of ''Thylacoleo carnifex'']]
+[[Image:Thylacoleo skull.jpg|thumb|Skull of ''Thylacoleo carnifex'']]
-Like other thylacoleonids, ''Thylacoleo'' had blade-like third [[premolar]] teeth in the upper and lower jaws, that functioned as the [[carnassial]] teeth, with these teeth being present much further forwards in the jaw than in other mammals.<ref name=":0">{{Cite journal |last=Werdelin |first=L |date=1988 |title=Circumventing a Constraint - the Case of Thylacoleo (Marsupialia, Thylacoleonidae) |url=http://www.publish.csiro.au/?paper=ZO9880565 |journal=Australian Journal of Zoology |language=en |volume=36 |issue=5 |pages=565 |doi=10.1071/ZO9880565 |issn=0004-959X}}</ref><ref name="rspb.royalsocietypublishing.org" /> Compared to earlier thyacoleonids, the third premolars were considerably larger.<ref name=":3">{{Cite journal |last=Gillespie |first=Anna K. |date=2023-01-16 |title=Two new marsupial lion taxa (Marsupialia, Thylacoleonidae) from the early and Middle Miocene of Australia |url=https://www.tandfonline.com/doi/full/10.1080/03115518.2022.2152096 |journal=Alcheringa: An Australasian Journal of Palaeontology |language=en |pages=1–16 |doi=10.1080/03115518.2022.2152096 |issn=0311-5518}}</ref>''Thylacoleo'' had a proportionally large pair of first [[incisor|incisors]] in the upper and lower jaws, which functioned analogously to other carnivores [[Canine tooth|canine teeth]].<ref name=":0" /><ref>{{cite journal |last1=Wells |first1=Roderick T. |last2=Murray |first2=Peter F. |last3=Bourne |first3=Steven J. |year=2009 |title=Pedal morphology of the marsupial lion ''Thylacoleo carnifex'' (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia |journal=Journal of Vertebrate Paleontology |volume=29 |issue=4 |pages=1335–1340 |bibcode=2009JVPal..29.1335W |doi=10.1671/039.029.0424 |s2cid=86460654}}</ref> They also had true canines but they served little purpose as they were stubby and not very sharp.<ref name=":1">{{Cite web |title=Discovery and Interpretation |url=http://www.naturalworlds.org/thylacoleo/discovery/discovering/discovering_tc_1.htm |access-date=7 May 2019 |website=Natural Worlds}}</ref> Compared to earlier thylacoleonids, the number of [[Molar (tooth)|molar]] teeth was reduced.<ref name=":3" />
+Like other thylacoleonids, ''Thylacoleo'' had blade-like third [[premolar]] teeth in the upper and lower jaws, that functioned as the [[carnassial]] teeth, with these teeth being present much further forwards in the jaw than in other mammals.<ref name=":0">{{Cite journal |last=Werdelin |first=L |date=1988 |title=Circumventing a Constraint - the Case of Thylacoleo (Marsupialia, Thylacoleonidae) |url=http://www.publish.csiro.au/?paper=ZO9880565 |journal=Australian Journal of Zoology |language=en |volume=36 |issue=5 |pages=565 |doi=10.1071/ZO9880565 |issn=0004-959X}}</ref><ref name="rspb.royalsocietypublishing.org" /> Compared to earlier thyacoleonids, the third premolars were considerably enlarged.<ref name=":3">{{Cite journal |last=Gillespie |first=Anna K. |date=2023-01-16 |title=Two new marsupial lion taxa (Marsupialia, Thylacoleonidae) from the early and Middle Miocene of Australia |url=https://www.tandfonline.com/doi/full/10.1080/03115518.2022.2152096 |journal=Alcheringa: An Australasian Journal of Palaeontology |volume=47 |issue=4 |language=en |pages=506–521 |doi=10.1080/03115518.2022.2152096 |bibcode=2023Alch...47..506G |s2cid=256157821 |issn=0311-5518}}</ref> ''Thylacoleo'' also had a proportionally large pair of first [[incisor]]s in the upper and lower jaws, which functioned analogously to other carnivores [[Canine tooth|canine teeth]].<ref name=":0" /><ref>{{cite journal |last1=Wells |first1=Roderick T. |last2=Murray |first2=Peter F. |last3=Bourne |first3=Steven J. |year=2009 |title=Pedal morphology of the marsupial lion ''Thylacoleo carnifex'' (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia |journal=Journal of Vertebrate Paleontology |volume=29 |issue=4 |pages=1335–1340 |bibcode=2009JVPal..29.1335W |doi=10.1671/039.029.0424 |s2cid=86460654}}</ref> They also had true canines but they served little purpose as they were stubby and not very sharp.<ref name=":1">{{Cite web |title=Discovery and Interpretation |url=http://www.naturalworlds.org/thylacoleo/discovery/discovering/discovering_tc_1.htm |access-date=7 May 2019 |website=Natural Worlds}}</ref> Compared to earlier thylacoleonids, the number of [[Molar (tooth)|molar]] teeth was reduced.<ref name=":3" />
-Pound for pound, ''T. carnifex'' had the strongest bite of any [[mammal]] species, living or extinct; a ''T. carnifex'' weighing {{convert|101|kg|abbr=on}} had a bite comparable to that of a 250&nbsp;kg African [[lion]], and research suggests that ''Thylacoleo'' could hunt and take prey much larger than itself.<ref name="rspb.royalsocietypublishing.org">{{Cite journal | doi=10.1098/rspb.2004.2986| pmid=15817436| pmc=1564077| title=Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa| journal=Proceedings of the Royal Society B: Biological Sciences| volume=272| issue=1563| pages=619–625| year=2005| last1=Wroe| first1=S.| last2=McHenry| first2=C.| last3=Thomason| first3=J.}}</ref> Larger animals that were likely prey include ''[[Diprotodon]]'' spp. and [[Protemnodon|giant kangaroos]]. It seems improbable that ''Thylacoleo'' could achieve as high a bite force as a modern-day lion; however, this might have been possible when taking into consideration the size of its brain and skull. Carnivores usually have rather large brains when compared to [[Herbivore|herbivorous]] marsupials, which lessens the amount of bone that can be devoted to enhancing bite force. ''Thylacoleo'' however, is thought to have had substantially stronger muscle attachments and therefore a smaller brain.<ref>{{Cite web|url=https://laelaps.wordpress.com/2007/08/31/thylacoleo-carnifex-ancient-australias-marsupial-lion/|title=Thylacoleo carnifex, ancient Australia's marsupial lion|last=Switek|first=Brian|date=31 August 2007|website=Laelaps|access-date=8 May 2019}}</ref>
+Pound for pound, ''T. carnifex'' had the strongest bite of any [[mammal]] species, living or extinct; a ''T.&nbsp;carnifex'' weighing {{convert|101|kg|abbr=on}} had a bite comparable to that of a 250&nbsp;kg African [[lion]], and research suggests that ''Thylacoleo'' could hunt and take prey much larger than itself.<ref name="rspb.royalsocietypublishing.org">{{Cite journal | doi=10.1098/rspb.2004.2986| pmid=15817436| pmc=1564077| title=Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa| journal=Proceedings of the Royal Society B: Biological Sciences| volume=272| issue=1563| pages=619–625| year=2005| last1=Wroe| first1=S.| last2=McHenry| first2=C.| last3=Thomason| first3=J.}}</ref> Larger animals that were likely prey include ''[[Diprotodon]]'' spp. and [[Protemnodon|giant kangaroos]]. It seems improbable that ''Thylacoleo'' could achieve as high a bite force as a modern-day lion; however, this might have been possible when taking into consideration the size of its brain and skull. Carnivores usually have rather large brains when compared to [[Herbivore|herbivorous]] marsupials, which lessens the amount of bone that can be devoted to enhancing bite force. ''Thylacoleo'' however, is thought to have had substantially stronger muscle attachments and therefore a smaller brain.<ref>{{Cite web|url=https://laelaps.wordpress.com/2007/08/31/thylacoleo-carnifex-ancient-australias-marsupial-lion/|title=Thylacoleo carnifex, ancient Australia's marsupial lion|last=Switek|first=Brian|date=31 August 2007|website=Laelaps|access-date=8 May 2019}}</ref> Some later studies questioned the ability of the canine teeth to deliver a killing bite.<ref name="figu2016">{{cite journal |last1=Figueirido |first1=Borja |last2=Martín-Serra |first2=Alberto |last3=Janis |first3=Christine M. |year=2016 |title=Ecomorphological determinations in the absence of living analogues: The predatory behavior of the marsupial lion (Thylacoleo carnifex) as revealed by elbow joint morphology |url=https://research-information.bris.ac.uk/ws/files/57326592/Figueirido_Martin_Serra_Janis.pdf |journal=Paleobiology |volume=42 |issue=3 |pages=508–531 |bibcode=2016Pbio...42..508F |doi=10.1017/pab.2015.55 |s2cid=87168573 |hdl=1983/1f3ac566-0b08-48fb-b27c-63fe4256c138}}</ref>
-Using [[3D modeling]] based on [[X-ray computed tomography]] scans,  marsupial lions were found to be unable to use the prolonged, suffocating bite typical of living big cats. They instead had an extremely efficient and unique bite; the incisors would have been used to stab at and pierce the flesh of their prey while the more specialised carnassials crushed the [[Vertebrate trachea|windpipe]], severed the [[spinal cord]], and lacerated the major [[blood vessel]]s such as the [[carotid artery]] and [[jugular vein]]. Compared to an African lion which may take 15 minutes to kill a large catch, the marsupial lion could kill a large animal in less than a minute. The skull was so specialized for big game that it was very inefficient at catching smaller animals, which possibly contributed to its extinction.<ref>[https://www.sciencedaily.com/releases/2008/01/080117093440.htm "Extinct Marsupial Lion Tops African Lion In Fight To Death"], ''[[Science Daily]]'', 17 January 2008.</ref><ref>{{cite web |url=http://www.theaustralian.com.au/news/health-science/marsupial-lion-was-fast-killer/story-e6frg8gf-1111115343423 |title=Marsupial lion was fast killer |archiveurl=https://web.archive.org/web/20090417020211/http://www.theaustralian.news.com.au/story/0,25197,23069545-30417,00.html |website=The Australian |date=18 January 2008 |archivedate=17 April 2009 |first1=Leigh |last1=Dayton}}</ref>
+Using [[3D modeling]] based on [[X-ray computed tomography]] scans,  marsupial lions were found to be unable to use the prolonged, suffocating bite typical of living big cats. They instead had an extremely efficient and unique bite; the incisors would have been used to stab at and pierce the flesh of their prey while the more specialised carnassials crushed the [[Vertebrate trachea|windpipe]], severed the [[spinal cord]], and lacerated the major [[blood vessel]]s such as the [[carotid artery]] and [[jugular vein]]. Compared to an African lion which may take 15 minutes to kill a large catch, the marsupial lion could kill a large animal in less than a minute.{{dubious|talk=Dubious claim|date=September 2024}} The skull was so specialized for big game that it was very inefficient at catching smaller animals, which possibly contributed to its extinction.<ref>[https://www.sciencedaily.com/releases/2008/01/080117093440.htm "Extinct Marsupial Lion Tops African Lion In Fight To Death"], ''[[Science Daily]]'', 17 January 2008.</ref><ref>{{cite web |url=http://www.theaustralian.com.au/news/health-science/marsupial-lion-was-fast-killer/story-e6frg8gf-1111115343423 |title=Marsupial lion was fast killer |archiveurl=https://web.archive.org/web/20090417020211/http://www.theaustralian.news.com.au/story/0,25197,23069545-30417,00.html |website=The Australian |date=18 January 2008 |archivedate=17 April 2009 |first1=Leigh |last1=Dayton}}</ref>
 ===Postcranium===
 [[File:Thylacoleo skeleton in Naracoorte Caves (cropped).jpg|left|thumb|Skeleton at Naracoorte Caves]]
+''Thylacoleo'' had highly mobile and powerful forelimbs used to grapple prey, with each [[Manus (anatomy)|manus]] having a single very large retractable hooked claw set on large semi-opposable thumbs, which are suggested to have been used deal a killing blow.<ref name="figu2016" />
-''Thylacoleo'' had extremely strong forelimbs, with retractable, cat-like claws, a trait previously unseen in marsupials. ''Thylacoleo'' also possessed enormous hooked claws set on large semi-opposable thumbs, which were used to capture and disembowel prey. The long muscular tail was similar to that of a kangaroo. Specialised tail bones called [[Chevron (anatomy)|chevrons]] allowed the animal to balance on its back legs, and freed the front legs for slashing and grasping.<ref>{{Cite web|url=https://www.pbs.org/wgbh/nova/bonediggers/thyl-nf.html|title=NOVA &#124; Bone Diggers &#124; Anatomy of Thylacoleo &#124; PBS|website=www.pbs.org}}</ref>
-Retractable claws would remain sharp by protecting them from being worn down on hard surfaces. The first digits ("[[thumbs]]") on each hand were semi-opposable and bore an enlarged claw. [[Palaeontologist]]s believe this would have been used to grapple its intended prey, as well as providing it with a sure footing on tree trunks and branches. The hind feet had four functional toes, the first digit being much reduced in size, but possessing a roughened pad similar to that of [[Phalangeriformes|possums]], which may have assisted with climbing. The discovery in 2005 of a specimen which included complete hind feet provided evidence that the marsupial lion exhibited [[syndactyly]] (fused second and third toes) like other diprotodonts.<ref>{{cite journal|author1= Wells, R.T.|author2=Murray, P.F.|author3=Bourne, S.J.|year=2009|title=Pedal morphology of the marsupial lion ''Thylacoleo carnifex'' (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia|journal=Journal of Vertebrate Paleontology|volume=29|issue=4|pages=1335–1340|doi=10.1671/039.029.0424 |bibcode=2009JVPal..29.1335W |s2cid=86460654}}</ref>
+The hind feet had four functional toes, the first digit being much reduced in size, but possessing a roughened pad similar to that of [[Phalangeriformes|possums]], which may have assisted with climbing. The discovery in 2005 of a specimen which included complete hind feet provided evidence that the marsupial lion exhibited [[syndactyly]] (fused second and third toes) like other diprotodonts.<ref>{{cite journal|author1= Wells, R.T.|author2=Murray, P.F.|author3=Bourne, S.J.|year=2009|title=Pedal morphology of the marsupial lion ''Thylacoleo carnifex'' (Diprotodontia: Thylacoleonidae) from the Pleistocene of Australia|journal=Journal of Vertebrate Paleontology|volume=29|issue=4|pages=1335–1340|doi=10.1671/039.029.0424 |bibcode=2009JVPal..29.1335W |s2cid=86460654}}</ref>
-Its strong forelimbs and retracting claws mean that ''Thylacoleo'' possibly climbed trees and perhaps carried carcasses to keep the kill for itself (similar to the [[leopard]] today).<ref name="Claw">{{Cite journal|title =  Behaviour of the Pleistocene marsupial lion deduced from claw marks in a southwestern Australian cave|journal = Scientific Reports|volume = 6|pages = 21372|last1=Arman |first1=Samuel D. |last2=Prideaux |first2=Gavin J. |date = 15 February 2016|doi = 10.1038/srep21372|pmid = 26876952|pmc = 4753435|bibcode = 2016NatSR...621372A|s2cid=3548956}}</ref> The climbing ability would have also helped them climb out of caves, which could therefore have been used as dens to rear their young.<ref name="Evans2018">{{cite journal|last1= Evans|first1=A. R.|last2= Wells|first2=R. T.|last3= Camens|first3=A. B.|title= New skeletal material sheds light on the palaeobiology of the Pleistocene marsupial carnivore, ''Thylacoleo carnifex''|journal=PLOS ONE|volume= 13|issue= 12|year= 2018|pages= e0208020|doi= 10.1371/journal.pone.0208020|pmid=30540785|pmc=6291118|bibcode=2018PLoSO..1308020W|doi-access=free}}</ref>
+Its strong forelimbs and retracting claws mean that ''Thylacoleo'' possibly climbed trees and perhaps carried carcasses to keep the kill for itself (similar to the [[leopard]] today).<ref name="Claw">{{Cite journal|title =  Behaviour of the Pleistocene marsupial lion deduced from claw marks in a southwestern Australian cave|journal = Scientific Reports|volume = 6|pages = 21372|last1=Arman |first1=Samuel D. |last2=Prideaux |first2=Gavin J. |date = 15 February 2016|doi = 10.1038/srep21372|pmid = 26876952|pmc = 4753435|bibcode = 2016NatSR...621372A|s2cid=3548956}}</ref> The climbing ability would have also helped them climb out of caves, which could therefore have been used as dens to rear their young.<ref name="Evans2018">{{cite journal|last1= Evans|first1=A. R.|last2= Wells|first2=R. T.|last3= Camens|first3=A. B.|title= New skeletal material sheds light on the palaeobiology of the Pleistocene marsupial carnivore, ''Thylacoleo carnifex''|journal=PLOS ONE|volume= 13|issue= 12|year= 2018|pages= e0208020|doi= 10.1371/journal.pone.0208020|pmid=30540785|pmc=6291118|bibcode=2018PLoSO..1308020W|doi-access=free}}</ref> Specialised tail bones called [[Chevron (anatomy)|chevrons]] strengthened the tail, likely allowing the animal to use it to prop itself up while rearing on its hind legs, which may have been done when climbing or attacking prey.<ref name="Evans2018" />
+The [[lumbar]] region is relatively rigid and straight, and suggests that the lower back was relatively inflexible.<ref name="Evans2018" />
-Due to its unique predatory morphology, some scientists have claimed ''Thylacoleo'' to be the most specialised mammalian carnivore of all time.<ref>{{Cite web|url=http://news.nationalgeographic.com/news/2004/03/0305_040305_TVsuperpredator.html|title=Extinct Australian "Lion" Was Big Biter, Expert Says<!-- Bot generated title -->}}</ref>
+==Ecology==
-==Behaviour and diet==
-[[File:Thylacoleo vs Diprotodon.jpg|thumb|Restoration of ''Thylacoleo'' attacking ''[[Diprotodon]]''.]]
+[[File:Marsupial lion (Thylacoleo carnifex).png|thumb|Life restoration of ''Thylacoleo carnifex'']]
+When ''Thylacoleo'' was first described by Richard Owen, he considered it to be a carnivore, based on the morphology of its skull and teeth.<ref name="Owen1859XVI" /> However other anatomists, such as [[William Henry Flower]] disagreed. Flower was the first to place ''Thylacoleo'' with the Diprotodonts, noting its skull and teeth to be laid out more like those of the [[koala]] and the [[wombat]], and suggested that it was more likely a herbivore. Owen did not disagree with Flower's placement of ''Thylacoleo'' with the Diprotodonts, but still maintained that it was a carnivore, despite its herbivorous ancestry.<ref name="wired2009">{{cite magazine | url=https://www.wired.com/2009/06/thylacoleo-herbivore-or-carnivore/?intcid=inline_amp | title=Thylacoleo: Herbivore or Carnivore? | magazine=Wired | last1=Switek | first1=Brian }}</ref> Owen found little support in his lifetime, despite the pointing out of ''Thylacoleo'''s retractable claws, something only found in mammalian carnivores,<ref name="Owen1883XV">{{cite journal|doi=10.1098/rstl.1883.0015|title=XV. On the affinities of thylacoleo |journal=Philosophical Transactions of the Royal Society of London |year=1883 |volume=174 |pages=575–582 |s2cid=111347165|issn=0261-0523|doi-access=free}}</ref> and its lack of any ability to chew plant material.<ref name="wired2009" />  In 1911, a study by Spencer and Walcott claimed that certain marks on the bones of megafauna had been made by ''Thylacoleo'', but according to Horton (1979) they were not sufficiently rigorous, resulting in their arguments being strongly challenged by later scholars, such as Anderson (1929), and later Gill (1951, 1952, 1954), thereby leaving the issue unresolved.<ref name="bite">{{Cite journal |last1=Horton |first1=D. R. |last2=Wright |first2=R. V. S. |year=1981 |title=Cuts on Lancefield Bones: Carnivorous Thylacoleo, Not Humans, the Cause |journal=Archaeology in Oceania |volume=16 |issue=2 |pages=73–80 |doi=10.1002/j.1834-4453.1981.tb00009.x |jstor=40386545}}</ref>
+Besides the most common hypothesis that it was an active predator, a variety of other theories existed in the late 19th to early 20th centuries as to the diet and feeding of ''Thylacoleo'', with hypotheses of it being a scavenger filling the ecological niche of [[hyenas]],<ref name="scavenger">{{cite journal |last1=De Vis |first1=C. W. |year=1883 |title=On tooth-marked bones of extinct marsupials |url=https://www.biodiversitylibrary.org/part/28646 |journal=Proceedings of the Linnean Society of New South Wales |volume=8 |pages=187–190 |doi=10.5962/bhl.part.28646 |doi-access=free}}</ref> being a specialist of crocodile eggs,<ref name="Claw" /> or even a melon-eater.<ref name="melons">{{cite journal |last1=Anderson |first1=C. |year=1929 |title=Palaeontological notes no. 1. Macropus titan Owen and Thylacoleo carnifex Owen |journal=Records of the Australian Museum |volume=17 |pages=35–49 |doi=10.3853/j.0067-1975.17.1929.752 |doi-access=free}}</ref> As late as 1954, doubts were still being raised as to whether it was actually a hypercarnivore.<ref name="figu2016" />
-===Diet===
-When ''Thylacoleo'' was first described by Richard Owen, he considered it to be a carnivore, based on the morphology of its skull and teeth.<ref name="Owen1859XVI" /> However other anatomists, such as [[William Henry Flower]] disagreed. Flower was the first to place ''Thylacoleo'' with the Diprotodonts, noting its skull and teeth to be laid out more like those of the [[koala]] and the [[wombat]], and suggested that it was more likely a herbivore. Owen did not disagree with Flower's placement of ''Thylacoleo'' with the Diprotodonts, but still maintained that it was a carnivore, despite its herbivorous ancestry.<ref name= "wired2009" >{{cite magazine | url=https://www.wired.com/2009/06/thylacoleo-herbivore-or-carnivore/?intcid=inline_amp | title=Thylacoleo: Herbivore or Carnivore? | magazine=Wired | last1=Switek | first1=Brian }}</ref> Owen found little support in his lifetime, despite the pointing out of ''Thylacoleo'''s retractable claws, something only found in mammalian carnivores,<ref name="Owen1883XV" >{{cite journal|doi=10.1098/rstl.1883.0015|title=XV. On the affinities of thylacoleo |journal=Philosophical Transactions of the Royal Society of London |year=1883 |volume=174 |pages=575–582 |s2cid=111347165|issn=0261-0523|doi-access=free}}</ref> and its lack of any ability to chew plant material.<ref name="wired2009" /> In 1911, a study by Spencer and Walcott claimed that certain marks on the bones of megafauna had been made by ''Thylacoleo'', but according to Horton (1979) they were not sufficiently rigorous, resulting in their arguments being strongly challenged by later scholars, such as Anderson (1929), and later Gill (1951, 1952, 1954), thereby leaving the issue unresolved.<ref name="bite" /> In 1981, another paper was published arguing that certain cuts to bones of large marsupials had been caused by ''Thylacoleo''. This paper by Horton and Wright was able to counter earlier arguments that such marks were the result of humans, largely by pointing out the presence of similar marks on the opposite side of many bones. They concluded that humans were extremely unlikely to have made the marks in question, but that if so "they had set out to produce only marks consistent with what ''Thylacoleo'' would produce".<ref name="bite" /> Since then, the academic consensus has emerged that ''Thylacoleo'' was a predator and a hypercarnivore.<ref name="figu2016" />
+In 1981, another paper was published arguing that certain cuts to bones of large marsupials had been caused by ''Thylacoleo''. This paper by Horton and Wright was able to counter earlier arguments that such marks were the result of humans, largely by pointing out the presence of similar marks on the opposite side of many bones. They concluded that humans were extremely unlikely to have made the marks in question, but that if so "they had set out to produce only marks consistent with what ''Thylacoleo'' would produce".<ref name="bite" /> Since then, the academic consensus has emerged that ''Thylacoleo'' was a predator and a hypercarnivore.<ref name="figu2016" />
-Besides the most common hypothesis that it was an active predator, a variety of other theories existed in the late 19th to early 20th centuries as to the diet and feeding of ''Thylacoleo'', with hypotheses of it being a scavenger filling the ecological niche of [[hyenas]],<ref name="scavenger" >{{cite journal|doi=10.5962/bhl.part.28646|title=On tooth-marked bones of extinct marsupials |year=1883 |last1=De Vis |first1=C. W. |journal=Proceedings of the Linnean Society of New South Wales |volume=8 |pages=187–190 |url=https://www.biodiversitylibrary.org/part/28646 |doi-access=free }}</ref> being a specialist of crocodile eggs,<ref name="Claw" /> or even a melon-eater.<ref name="melons" >{{cite journal|doi=10.3853/j.0067-1975.17.1929.752|title=Palaeontological notes no. 1. Macropus titan Owen and Thylacoleo carnifex Owen |year=1929 |last1=Anderson |first1=C. |journal=Records of the Australian Museum |volume=17 |pages=35–49 |doi-access=free }}</ref> As late as 1954, doubts were still being raised as to whether it was actually a hypercarnivore.<ref name="figu2016" >{{cite journal|doi=10.1017/pab.2015.55|title=Ecomorphological determinations in the absence of living analogues: The predatory behavior of the marsupial lion (Thylacoleo carnifex) as revealed by elbow joint morphology |year=2016 |last1=Figueirido |first1=Borja |last2=Martín-Serra |first2=Alberto |last3=Janis |first3=Christine M. |journal=Paleobiology |volume=42 |issue=3 |pages=508–531 |bibcode=2016Pbio...42..508F |s2cid=87168573 |url=https://research-information.bris.ac.uk/ws/files/57326592/Figueirido_Martin_Serra_Janis.pdf }}</ref>
+The marsupial lion's limb proportions and muscle mass distribution indicate that, although it was a powerful animal, it was not a particularly fast runner. Paleontologists conjecture that it was an [[ambush predator]], possibly using leaping.<ref name="Evans2018" /> Incisions on bones of the extinct large kangaroo ''[[Macropus titan]]'', and the general morphology of ''Thylacoleo'' suggests that it fed in a similar manner to modern cheetahs, by using their sharp teeth to slice open the ribcage of their prey, thereby accessing the internal organs. They may have killed by using their front claws as either stabbing weapons or as a way to grab their prey with strangulation or suffocation.<ref name="bite" />
-The marsupial lion's limb proportions and muscle mass distribution indicate that, although it was a powerful animal, it was not a particularly fast runner. Paleontologists conjecture that it was an [[ambush predator]], either sneaking up and then leaping upon its prey, or dropping down on it from overhanging tree branches. This is consistent with the depictions of the animal as striped: camouflage of that kind is needed for stalking and hiding in a largely forested habitat (like [[tigers]]) rather than chasing across open spaces (like [[lions]]).<ref name = guard>{{cite news|title='Like a demon in a medieval book': is this how the marsupial lion killed prey?|url=https://www.theguardian.com/environment/georgemonbiot/2014/apr/03/australia-marsupial-lion-kangaroos-prey|accessdate=22 October 2014|newspaper=The Guardian|date=2014-04-03|last1=Monbiot|first1=George}}</ref> Trace fossils in the form of claw marks and bones from caves in [[Western Australia]] analyzed by Gavin Prideaux et al. indicate marsupial lions could also climb rock faces, and likely reared their young in such caves as a way of protecting them from potential predators.<ref>{{Cite news | url=https://www.bbc.com/news/world-australia-35557269 | title=Marsupial lion 'could climb trees'| work=BBC News| date=2016-02-15}}</ref> It is thought to have hunted large animals such as the enormous ''[[Diprotodon]]'' and giant [[browsing (predation)|browsing]] [[kangaroo]]s like ''[[Sthenurus]]'' and ''[[Procoptodon]]'', and competed with other predatory animals such as the giant monitor lizard, ''[[Megalania]]'', and terrestrial crocodiles such as ''[[Quinkana]]''.  The marsupial lion may have cached kills in trees in a manner similar to the modern [[leopard]].<ref>[http://www.museum.wa.gov.au/exhibitions/online/thylacoleo/hunter.asp Western Australian Museum Thylacoleo (panel 3)] {{webarchive|url=https://web.archive.org/web/20080727131817/https://www.museum.wa.gov.au/exhibitions/online/thylacoleo/hunter.asp |date=2008-07-27 }} at Western Australian Museum</ref> Like many predators, it was probably also an opportunistic [[scavenger]], feeding on [[carrion]] and driving off less powerful predators from their kills. It also may have shared behaviours exhibited by recent [[diprotodont]] marsupials such as kangaroos, like digging shallow holes under trees to reduce body temperature during the day.<ref>{{cite book |last1=Tyndale-Biscoe |first1=Hugh |title=Life of marsupials |date=2005 |publisher=CSIRO |location=Collingwood, Vic. |isbn=978-0-643-09220-4 }}{{page needed|date=August 2018}}</ref>
+Like many predators, it was probably also an opportunistic [[scavenger]], feeding on [[carrion]] and driving off less powerful predators from their kills. It also may have shared behaviours exhibited by recent [[diprotodont]] marsupials such as kangaroos, like digging shallow holes under trees to reduce body temperature during the day.<ref>{{cite book |last1=Tyndale-Biscoe |first1=Hugh |title=Life of marsupials |date=2005 |publisher=CSIRO |isbn=978-0-643-09220-4 |location=Collingwood, Vic.}}{{page needed|date=August 2018}}</ref>
-===Senses===
-[[Image:Thylacoleo BW.jpg|thumb|Restoration of ''T. carnifex'']]
-[[Computed tomography|CT]] scans of a well-preserved [[skull]] have allowed scientists to study internal structures and create a brain [[endocast]] showing the surface features of the animal's brain. The [[parietal lobe]]s, [[visual cortex]], and [[olfactory bulb]]s of the [[cerebrum]] were enlarged, indicating the marsupial lion had good senses of hearing, sight, and smell, as might be expected of an active predator. Also, a pair of blind canals within the [[nasal cavity]] were probably associated with detecting [[pheromone]]s as in the [[Tasmanian devil]]. This indicates it most likely had seasonal mating habits and would "sniff out" a mate when in season.<ref name="abc.net.au">[http://www.abc.net.au/catalyst/stories/s1717424.htm Thylacoleo "The Beast of the Nullarbor"], ''[[Catalyst (TV series)|Catalyst]]'', Western Australian Museum, Storyteller Media Group and ABC TV, 17 August 2006.</ref>
+Trace fossils in the form of claw marks and bones from caves in [[Western Australia]] analyzed by Gavin Prideaux et al. indicate marsupial lions could also climb rock faces, and likely reared their young in such caves as a way of protecting them from potential predators.<ref>{{Cite news | url=https://www.bbc.com/news/world-australia-35557269 | title=Marsupial lion 'could climb trees'| work=BBC News| date=2016-02-15}}</ref>
-==Feeding==
-The feeding behaviour of ''Thylacoleo'' remains a topic of academic debate, largely due the lack of any living analogue.<ref name="figu2016" />
-While considered a powerful hunter, and a fierce predator, it has been theorized that due to its physiology ''Thylacoleo'' was, in fact, a slow runner, limiting its ability to chase prey. Analysis of its scapula suggests "walking and trotting, rather than climbing ... the pelvis similarly agrees with that of ambulators and cursors [walkers and runners]". These bones indicate that ''Thylacoleo'' was a slow to medium-paced runner, which is likely to mean it was an ambush predator. That fits with the stripes: camouflage of the kind one would need for stalking and hiding in a largely forested habitat (like tigers or leopards) rather than chasing across open spaces (like lions)."<ref>{{Cite news|url=https://www.theguardian.com/environment/georgemonbiot/2014/apr/03/australia-marsupial-lion-kangaroos-prey/ | title='Like a demon in a medieval book': Is this how the marsupial lion killed prey?| newspaper=The Guardian| date=2014-04-03| last1=Monbiot| first1=George}}</ref> It may have functioned generally much like a larger analog of the [[Sarcophilus harrisii|Tasmanian devil]].<ref name="Evans2018" /> New evidence also suggests that it may have been arboreal, and was at the very least capable of climbing trees.<ref name="Claw" />
+Analysis of finds on the Nullabor Plain suggests that ''Thylacoleo carnifex'' inhabited open, arid environments similar to those found across much of Australia today.<ref>{{Cite journal |last1=Prideaux |first1=Gavin J. |last2=Long |first2=John A. |last3=Ayliffe |first3=Linda K. |last4=Hellstrom |first4=John C. |last5=Pillans |first5=Brad |last6=Boles |first6=Walter E. |last7=Hutchinson |first7=Mark N. |last8=Roberts |first8=Richard G. |last9=Cupper |first9=Matthew L. |last10=Arnold |first10=Lee J. |last11=Devine |first11=Paul D. |last12=Warburton |first12=Natalie M. |date=January 2007 |title=An arid-adapted middle Pleistocene vertebrate fauna from south-central Australia |url=https://www.nature.com/articles/nature05471 |journal=Nature |language=en |volume=445 |issue=7126 |pages=422–425 |doi=10.1038/nature05471 |pmid=17251978 |bibcode=2007Natur.445..422P |issn=0028-0836}}</ref>
-Incisions on bones of ''Macropus titan'', and the general morphology of ''Thylacoleo'' suggests that it fed in a similar manner to modern cheetahs, by using their sharp teeth to slice open the ribcage of their prey, thereby accessing the internal organs. They may have killed by using their front claws as either stabbing weapons or as a way to grab their prey with strangulation or suffocation.<ref name="bite">{{Cite journal|title =Cuts on Lancefield Bones: Carnivorous Thylacoleo, Not Humans, the Cause  |jstor=40386545| volume=16|issue=2|pages=73–80|journal= Archaeology in Oceania |last1=Horton|first1=D. R.|last2=Wright|first2=R. V. S.|year=1981|doi=10.1002/j.1834-4453.1981.tb00009.x}}</ref>
+A 1985 study suggested that ''Thylacoleo carnifex'' was an apex predator that primarily fed on large bodied prey, which may have included the large kangaroos ''[[Sthenurus]]'', ''[[Procoptodon]], [[Protemnodon]], [[Macropus]]'' and ''[[Osphranter|Ostphranter]]'' as well as possibly the largest Australian marsupial, the rhinoceros-sized vombatoid ''[[Diprotodon]]''.<ref>{{Cite journal |last=Case |first=J. A. |date=1985-01-01 |title=Differences in prey utilization by Pleistocene marsupial carnivores, Thylacoleo carnifex (Thylacoleonidae) and Thylacinus cynocephalus (Thylacinidae). |url=https://www.publish.csiro.au/AM/AM85002 |journal=Australian Mammalogy |language=en |volume=8 |issue=1 |pages=45–52 |doi=10.1071/AM85002 |issn=0310-0049}}</ref>
-==Palaeoecology==
+== Extinction ==
-[[File:Skull of the Marsupial Lion (Thylacoleo carnifex).jpg|thumb|Mounted display of ''Thylacoleo carnifex'' at the Wonambi Fossil Centre, Naracoorte Caves National Park, South Australia.]]
+{{Main|Late Pleistocene extinctions}}
-Numerous fossil discoveries indicate the marsupial lion was distributed across much of the [[Australia]]n [[continent]]. A large proportion of its environment would have been similar to the southern third of Australia today, [[semiarid]], open [[scrubland|scrub]] and [[woodland]] punctuated by [[Depression (geology)|waterhole]]s and water courses.{{Citation needed|date=June 2008}}
+''Thylacoleo'' is thought to have become extinct around 40,000 years ago as part of the [[Quaternary extinction event|Late Pleistocene megafauna extinctions]], essentially simultanteously with the vast majority of [[Australian megafauna]]. It has been contested as to the relative importance of climatic change vs the impact of recently arrived [[Indigenous Australians]] (who arrived in Australia around 50-60,000 years ago) in the extinctions. There is limited evidence of human interaction with extinct megafauna in Australia.<ref>{{Cite journal |last1=Hocknull |first1=Scott A. |last2=Lewis |first2=Richard |last3=Arnold |first3=Lee J. |last4=Pietsch |first4=Tim |last5=Joannes-Boyau |first5=Renaud |last6=Price |first6=Gilbert J. |last7=Moss |first7=Patrick |last8=Wood |first8=Rachel |last9=Dosseto |first9=Anthony |last10=Louys |first10=Julien |last11=Olley |first11=Jon |last12=Lawrence |first12=Rochelle A. |date=2020-05-18 |title=Extinction of eastern Sahul megafauna coincides with sustained environmental deterioration |journal=Nature Communications |language=en |volume=11 |issue=1 |page=2250 |doi=10.1038/s41467-020-15785-w |issn=2041-1723 |pmc=7231803 |pmid=32418985|bibcode=2020NatCo..11.2250H }}</ref>
+== See also ==
-It would have coexisted with many of the so-called [[Australian megafauna]] such as ''Diprotodon'', giant kangaroos, and ''Megalania'', as well as giant [[wallaby|wallabies]] like ''[[Protemnodon]]'', the giant [[wombat]] ''[[Phascolonus]]'', the giant snake ''[[Wonambi]]'', and the [[Dromornithidae|thunderbird]] ''[[Genyornis]]''.<ref name="abc.net.au"/> ''T. hilli'' was a similar size to a contemporaneous thylacoleonid species, ''[[Wakaleo alcootaensis]]'', and may have occupied habitat to the exclusion of that carnivore.<ref name="Murray1990">{{cite journal |last1=Murray |first1=P. |last2=Megirian |first2=D. |title=Further observations on the morphology of Wakaleo vanderleueri (Marsupialia:Thylacoleonidae) from the mid-Miocene Camfield beds, Northern Territory |journal=The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences |date=1990 |volume=7 |issue=1 |pages=91–102 |url=https://www.biodiversitylibrary.org/part/262795}}</ref>
-Australia's [[Pleistocene megafauna]] would have been the prey for the agile ''T. carnifex'', who was especially [[adaptation|adapted]] for hunting large animals, but was not particularly suited to catching smaller prey. The relatively quick reduction in the numbers of its primary food source around 40,000 to 50,000 years ago probably led to the decline and eventual extinction of the marsupial lion. The arrival of [[human]]s in Australia and the use of [[fire-stick farming]] precipitated their decline.<ref name=bbc120324>{{cite news | url = https://www.bbc.co.uk/news/science-environment-17488447 | title = 'Humans killed off Australia's giant beasts' | publisher = [[BBC News]] | date = 24 March 2012 }}</ref> The extinction of ''T. carnifex'' makes [[Australia]] unique from the other continents because no substantial, [[Apex predators|apex mammalian predators]] have replaced the marsupial lions after their disappearance.<ref name="Academic Search Premier">{{cite journal |doi=10.1111/j.1461-0248.2009.01347.x |pmid=19614756 |title=Predator interactions, mesopredator release and biodiversity conservation |journal=Ecology Letters |volume=12 |issue=9 |pages=982–98 |year=2009 |last1=Ritchie |first1=Euan G |last2=Johnson |first2=Christopher N |doi-access=free |hdl=10536/DRO/DU:30039763 |hdl-access=free }}</ref>
-===Extinction===
-[[File:Collections of the Australian Museum - 7.jpg|right|thumb|220px|''Thylacoleo carnifex'' sculpture at the Australian Museum]]
-It was believed{{by whom|date=March 2022}} that the extinction was due to the climate changes, but human activities as an extinction driver of the most recent species is possible yet unproven. There is a growing consensus{{by whom|date=October 2022}} that the extinction of the [[megafauna]] was caused by progressive drying starting about 700,000 years ago (700 ka).{{citation needed|date=October 2022}} It is revealed recently{{when|date=October 2022}} that there was a major change in glacial-[[interglacial]] cycles after ~450 ka. As for human involvement's contribution to the extinction, one argument is that the arrival of humans was coincident with the disappearance of all the extinct megafauna. This is supported by the claims that during [[Marine isotope stage|MIS3]], climatic conditions are relatively stable and no major climate change would cause the mass extinction of megafauna including ''Thylacoleo''.<ref>{{Cite web|url = http://www.pnas.org/content/110/22/8777.full.pdf|title = Climate change frames debate over the extinction of megafauna in Sahul (Pleistocene Australia-New Guinea)}}</ref>
-Although believed to have been a victim of climate change, some scientists{{which|date=October 2022}} now believe ''Thylacoleo'' to have been exterminated by humans altering the ecosystem with fire in addition to hunting its prey. "They found ''Sporormiella'' spores, which grow in herbivore dung, virtually disappeared around 41,000 years ago, a time when no known climate transformation was taking place. At the same time, the incidence of fire increased, as shown by a steep rise in charcoal fragments. It appears that humans, who arrived in Australia around this time, hunted the megafauna to extinction".<ref>{{Cite news|url=https://www.bbc.com/news/science-environment-17488447|title=Humans killed off Australia's giant beasts|date=24 March 2012|work=BBC|access-date=7 May 2019|language=en-GB}}</ref> Following the extinction of ''T. carnifex'', no other [[Apex predator|apex mammalian predators]] have taken its place.<ref>{{Cite journal|title = Predator interactions, mesopredator release and biodiversity conservation|journal = Ecology Letters|date = 2009-09-01|issn = 1461-0248|pages = 982–998|volume = 12|issue = 9|doi = 10.1111/j.1461-0248.2009.01347.x|first1 = Euan G.|last1 = Ritchie|first2 = Christopher N.|last2 = Johnson|pmid=19614756|doi-access = free|hdl = 10536/DRO/DU:30039763|hdl-access = free}}</ref>
-==Discoveries==
-[[Image:Thylacoskullcope.jpg|thumb|Drawing of ''T. carnifex'' skull fragments, 1859]]
-The first specimens of ''Thylacoleo'' were collected in the early 1830s from the Wellington Valley region of New South Wales by Major (later Sir) Thomas Mitchell, however they were not recognised as ''Thylacoleo'' at the time.<ref name=":1" />
-The species was first described by [[Richard Owen]] in 1859,<ref name=":1" /> from a fragmentary specimen discovered by William Adeney near Lake Colungolac, near Camperdown in [[Victoria, Australia|Victoria]]<ref name = "Owen1859XVI">{{cite journal|doi=10.1098/rstl.1859.0016|title=XVI. On the fossil mammals of Australia.— Part I. Description of a mutilated skull of a large marsupial carnivore (Thylacoleo carnifex, Owen), from a calcareous conglomerate stratum, eighty miles S. W. Of Melbourne, Victoria |journal=Philosophical Transactions of the Royal Society of London |year=1859 |volume=149 |pages=309–322 |s2cid=110651400 }}</ref>
-In 2002, eight remarkably complete skeletons of ''T. carnifex'' were discovered in a limestone cave under [[Nullarbor Plain]], where the animals fell through a narrow opening in the plain above. Based on the placement of their skeletons, at least some survived the fall, only to die of thirst and starvation.<ref>{{cite web|url=http://news.bbc.co.uk/2/hi/science/nature/6296029.stm |website=BBC News|title=Caverns give up huge fossil haul|date=25 January 2007}}</ref><ref>{{Cite web |url=http://news.nationalgeographic.com/news/2002/08/0807_020731_TVmegafauna.html |title=Australian Cave Yields Giant Animal Fossils |access-date=2009-09-03 |archive-date=2009-05-24 |archive-url=https://web.archive.org/web/20090524120938/http://news.nationalgeographic.com/news/2002/08/0807_020731_TVmegafauna.html |url-status=dead }}</ref>
-[[Image:Thylacoleo carnifex cave art.jpg|thumb|Tracing of [[cave art]] from the [[Kimberley (Western Australia)|Kimberley]] discovered in 2008, which was speculated to represent a drawing of the marsupial lion, but is more likely to be a drawing of a [[thylacine]]]]
-In 2008, rock art depicting what some speculate to be ''Thylacoleo'' was discovered on the northwestern coast of the Kimberley. However, the [[thylacine]], a marsupial that had a striped coat like depicted in the rock art, has been argued to be the more likely subject of the work.<ref>{{Cite journal |last=Welch |first=David M. |date=June 2015 |title=Thy Thylacoleo is a thylacine |url=http://www.tandfonline.com/doi/full/10.1080/03122417.2015.11682043 |journal=Australian Archaeology |language=en |volume=80 |issue=1 |pages=40–47 |doi=10.1080/03122417.2015.11682043 |s2cid=146358676 |issn=0312-2417}}</ref> The drawing represented only the second example of megafauna depicted by the indigenous inhabitants of Australia. The image contains details that would otherwise have remained only conjecture; the tail is depicted with a tufted tip, it has pointed ears rather than rounded, and the coat is striped. The prominence of the eye, a feature rarely shown in other animal images of the region, raises the possibility that the creature may have been a [[Nocturnality|nocturnal]] hunter.<ref name="Willing">{{cite web | last1 = Akerman | first1 = Kim | last2  = Willing | first2 = Tim| date = March 2009 | title = An ancient rock painting of a marsupial lion, ''Thylacoleo carnifex'', from the Kimberley, Western Australia | work = [[Antiquity (journal)|Antiquity]]| url = http://www.antiquity.ac.uk/projgall/akerman319/ | access-date = 11 December 2012 }}</ref> In 2009, a second image was found that depicts a ''Thylacoleo'' interacting with a hunter who is in the act of spearing or fending the animal off with a multiple-barbed spear. Much smaller and less detailed than the 2008 find, it may depict a [[thylacine]], but the comparative size indicates a ''Thylacoleo'' is more likely, meaning that it is possible that ''Thylacoleo'' was extant until more recently than previously thought.<ref name="Akerman">{{cite web | last = Akerman | first = Kim | date = December 2009 | title = Interaction between humans and megafauna depicted in Australian rock art? | work = [[Antiquity (journal)|Antiquity]] | url = http://www.antiquity.ac.uk/projgall/akerman322/ | access-date = 11 December 2012 }}</ref>
-In 2016, [[trace fossils]] in Tight Entrance Cave were identified as being the scratch marks of a ''Thylacoleo''.<ref name="Claw" />
-==Fossils==
-The first ''Thylacoleo'' fossil findings, discovered by [[Thomas Mitchell (explorer)|Thomas Mitchell]] were found in the 1830s in the Wellington Valley of New South Wales, though not recognised as such at the time. The generic holotype, consisting of broken teeth, jaws, and a skull was discovered by a pastoralist, William Avery, near Lake Colungolac from which the species ''Thylacoleo carnifex'' was described by [[Richard Owen]].<ref name="Owen1859XVI" /> It was not until 1966 that the first nearly-complete skeleton was found. The only pieces missing were a foot and the tail. Currently, the [[Nullarbor Plain]] of West Australia remains to be the greatest finding site. These fossils now reside at the Australian Museum.<ref>{{Cite web|url=http://australianmuseum.net.au/Thylacoleo-carnifex|title=Thylacoleo carnifex|last=Musser|first=Anne|date=29 November 2018|website=Australian Museum|access-date=7 May 2019}}</ref><ref name="Owen1883XV" />
-It was reported that in 2012, an accumulation of vertebrate trace and body fossils were found in the Victorian Volcanic Plains in southeastern Australia. It was determined that ''Thylacoleo'' was the only taxon that represented three divergent fossil records: skeletal, footprints, and bite marks. What this suggests is that these large carnivores had behavioral characteristics that could have increased their likelihood of their presence being detected within a fossil fauna.<ref>{{Cite journal|title = Contemporaneous Trace and Body Fossils from a Late Pleistocene Lakebed in Victoria, Australia, Allow Assessment of Bias in the Fossil Record|journal = PLOS ONE|date = 2013-01-02|pmc = 3534647|pmid = 23301008|pages = e52957|volume = 8|issue = 1|doi = 10.1371/journal.pone.0052957|first1 = Aaron Bruce|last1 = Camens|first2 = Stephen Paul|last2 = Carey|bibcode = 2013PLoSO...852957C|doi-access = free}}</ref>
-A characteristic seen in the remains of skull fragments is a set of carnassial teeth, suggesting the carnivorous habits of ''Thylacoleo''. Tooth fossils of the ''Thylacoleo'' exhibit specific degrees of erosion that are credited to the utility of the carnassial teeth remains as they were used for hunting and consuming prey in a prehistoric Australia teeming with other megafauna. The specialisation found in the dental history of the marsupial indicates its status in the predatory hierarchy in which it existed.<ref name= "Owen1866II">{{cite journal|doi=10.1098/rstl.1866.0004|title=IV. On the fossil mammals of Australia. - Part II. Description of an almost entire skull of the thylacoleo carnifex, Owen, from a freshwater deposit, Darling Downs, Queensland |journal=Philosophical Transactions of the Royal Society of London |year=1866 |volume=156 |pages=73–82 |s2cid=110169146 |doi-access=free }}</ref>
+* [[Drop bear]]
 {{clear}}
@@ Line 119: / Line 91: @@
 ==External links==
 {{Wikispecies|Thylacoleo}}
-* [http://phys.org/news/2013-05-evidence-theory-humans-megafauna.html New study finds no evidence for theory humans wiped out megafauna]
+* [https://phys.org/news/2013-05-evidence-theory-humans-megafauna.html New study finds no evidence for theory humans wiped out megafauna]
 * [http://www.naturalworlds.org/thylacoleo/index.htm ''Thylacoleo'' - Australia's Marsupial Lion]
 * [https://web.archive.org/web/20101124102953/http://thylacoleo.com/thylacoleo.php ''Thylacoleo'' in Pleistocene Australia]
 * [https://web.archive.org/web/20070426142324/http://www.bio.usyd.edu.au/staff/swroe/swroe.htm Steve Wroe's Web Page: Australian Megafauna]
 * [https://web.archive.org/web/20080727131817/https://www.museum.wa.gov.au/exhibitions/online/thylacoleo/hunter.asp Western Australian Museum: ''Thylacoleo'' - a voracious hunter]
-* [http://www.samuseum.sa.gov.au/Journals/RSAM/RSAM_v017/rsam_v017_p277p284.pdf  PLEDGE. N 1977, A NEW SPECIES OF ''THYLACOLEO'' (MARSUPIALIA: THYLACOLEONIDAE) WITH NOTES ON THE OCCURRENCES AND DISTRIBUTION OF THYLACOLEONIDAE IN SOUTH AUSTRALIA]{{Dead link|date=June 2018 |bot=InternetArchiveBot |fix-attempted=no }}
-{{Taxonbar|from=Q312099}}
+{{Vombatiformes}}{{Taxonbar|from=Q312099}}
 [[Category:Prehistoric vombatiforms]]