Temporal range: Early – Late Pleistocene
The marsupial lion (Thylacoleo carnifex) is an extinct species of carnivorous marsupial mammal that lived in Australia from the early to the late Pleistocene (1,600,000–46,000 years ago). Despite its name, it is not closely related to the lion, but is a member of the order Diprotodontia.
The marsupial lion is the largest meat-eating mammal known to have ever existed in Australia, and one of the largest marsupial carnivores from anywhere in the world (although see Thylacosmilus and Borhyaena). 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 100 to 130 kg (220 to 290 lb) in weight, although individuals heavier than 160 kg (350 lb) might not have been uncommon. This would make it quite comparable to female lions and tigers in general size.
The animal was extremely robust with powerfully built jaws and very strong fore limbs. It possessed retractable claws, a unique trait among marsupials. This would have allowed the claws to remain sharp by protecting them from being worn down on hard surfaces. The claws were well-suited to securing prey and for climbing trees. The first digits ("thumbs") on each hand were semiopposable and bore an enlarged claw. Palaeontologists 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 possums, which may have assisted with climbing. It is unclear whether the marsupial lion exhibited syndactyly (fused second and third toes) like other diprotodonts.
The marsupial lion's hindquarters were also well-developed, although to a lesser extent than the front of the animal. Remains of the animal show it had a relatively thick and strong tail and the vertebrae possessed chevrons on their undersides where the tail would have contacted the ground. These would have served to protect critical elements such as nerves and blood vessels if the animal used its tail to support itself when on its hind legs, much like present day kangaroos do. Taking this stance would free up its fore limbs to tackle or slash at its intended victim.
The ancestors of thylacoleonids are believed to have been herbivores, which is unusual for carnivores but cranial features and arboreal characteristics suggest a vombatoidea evolutionary descent. Possum-like features were once thought to indicate that the marsupial lion's evolutionary path was from a phalangeriform ancestor, however, scientists agree that more prominent features suggest a vombatiform ancestory; today represented by Lasiorhinus krefftii, Lasiorhinus latifrons, Phascolarctos cinereus, and Vombatus ursinus.
The marsupial lion was a highly specialised carnivore, as is reflected in its dentition. Like other diprotodonts, it possessed enlarged incisors on both the upper (maxillae) and lower (mandibles) jaws. These teeth (the lower in particular) were shaped much more like the pointed canine teeth of animals such as dogs and cats than those of kangaroos. The most unusual feature of the creature's dentition were the huge, blade-like carnassial premolars on either side of its jaws. The top and bottom carnassials worked together like shears and would have been very effective at slicing off chunks of flesh from carcasses and cutting through bone.
The jaw muscle of the marsupial lion was exceptionally large for its size, giving it an extremely powerful bite. Biometric calculations show, considering size, it had the strongest bite of any known mammal, living or extinct; a 100-kg individual would have had a bite comparable to that of a 250-kg African lion. 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. The skull was so specialized for big game, it was very inefficient at catching smaller animals, which possibly contributed to its extinction.
The marsupial lion's limb proportions and muscle mass distribution indicate, although it was a powerful animal, it was not a particularly fast runner. Palaeontologists 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. It is thought to have hunted large animals such as the enormous Diprotodon and giant browsing kangaroos like Sthenurus and Procoptodon. Although it may have shared at least part of its range with the 6-m-long megalania, the marsupial lion remains Australia's most ferocious apex predator because it possessed the most powerful bite than any other animal in its weight class. The marsupial lion may have cached kills in trees in a manner similar to the modern leopard. Like many predators, it was probably also an opportunistic scavenger, feeding on carrion and driving off less powerful predators from their kills.
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 lobes, visual cortex, and olfactory bulbs 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 pheromones as in the Tasmanian devil. This indicates it most likely had seasonal mating habits and would "sniff out" a mate when in season.
Numerous fossil discoveries indicate the marsupial lion was distributed across much of the Australian continent. A large proportion of its environment would have been similar to the southern third of Australia today, semiarid, open scrub and woodland punctuated by waterholes and water courses.
It would have coexisted with many of the so-called Australian megafauna such as the previously mentioned Diprotodon, giant kangaroos, and Megalania, as well as giant wallabies like Protemnodon, the giant wombat Phascolonus, and the thunderbird Genyornis.
Australia's Pleistocene megafauna would have been the prey for the agile Thylacoleo carnifex, who was especially 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 humans in Australia and the use of fire-stick farming precipitated their decline. The extinction of Thylaceleo carnifex makes Australia unique from the other continents because no substantial, apex mammalian predators have replaced the marsupial lions after their disappearance.
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 charcteristics led Thylaceleo 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. 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 metric tons. 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. 
Fossils of the marsupial lion have been found at several sites in Australia since the mid-19th century. A complete articulated skeleton was discovered in limestone caves under the Nullarbor Plain in 2002. The ends of the limb bones were not fully fused, indicating the animal was not full-grown.
Unlike most fossils, these bones were not mineralised and had been preserved in this state for about 500,000 years by the low humidity and cool temperature of the cave. The partial remains of 10 other individuals were found in this or nearby caves, along with hundreds of other specimens of other animals.
The animals apparently fell to their deaths tens of metres below, through narrow openings in the roof of the caves known as sinkholes. The caves and sinkholes were formed by groundwater slowly dissolving and eroding the limestone forming the bed of the plain (once a shallow sea).
Possible marsupial lion trace fossils have been found in a lake bed in south-western Victoria, along with trackways of a vombatid, the diprotodontid Diprotodon optatum, and a macropodid. The footprints were imprinted over a short period of time which may suggest an association between the marsupial lion and the other taxa present.  In addition, marsupial lion body fossils have been found in the same area and are dated around the same time as its trace fossils, a coincidence that is extremely rare and that may aid in a more complete assessment of the biodiversity in Australia during the Pleistocene epoch. Other fossils found at the site have bite marks that were presumably caused by the marsupial lion. 
Studies research suggest that most of mega fauna animals can survive and adapt to extreme weather climate change, therefore it is unlikely to be the cause of the extinction of the Thylacoleo carnifex. With that being said, scientists are now investigating the next possible cause which is the arrival of humans to Australia and how their actions could be the cause of the extinction of the Thylacoleo carnifex.
When humans came to Australia, they probably burnt the landscape of Australia to enhance their survival.  Changing the ecosystem of the once grassy woodland to desert shrub, the humans' actions may result in the extinction of plants that large and small animals feed off of. As a result, humans reduced the number of prey that big carnivores depend on (carnivores are sensitive to prey abundance). If humans reduce the number of potential prey below a threshold then large predators like the Thylacoleo carnifex will likely starve to death.
- Thylacoleo (panel 1) at Western Australian Museum
- 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): 489–498. doi:10.1071/ZO99006.
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- Thylacoleo carnifex at Australian Museum
- 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): 619–25. doi:10.1098/rspb.2004.2986. PMC 1564077. PMID 15817436.
- "Extinct Marsupial Lion Tops African Lion In Fight To Death", Science Daily, 17 January 2008.
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- Western Australian Museum Thylacoleo (panel 3) at Western Australian Museum
- Thylacoleo "The Beast of the Nullarbor", Catalyst, Western Australian Museum, Storyteller Media Group and ABC TV, 17 August 2006.
- "'Humans killed off Australia's giant beasts'". BBC News. 24 March 2012.
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- Black, Karen; Price, Gilbert; Archer, Michael; Hand, Suzanne (2014). "Bearing up well? Understanding the past, present and future of Australia's koalas". Gondwana Research 25 (3): 1186–1201. Retrieved 17 October 2014.
- Amalfi, Carmelo (31 July 2002). "Beneath the desert, the past blooms". The Age (Melbourne). Retrieved 2012-09-13.
- Carey, Stephen P.; Camens, Aaron B.; Cupper, Matthew L.; Grün, Rainer; Hellstrom, John C.; McKnight, Stafford W.; Mclennan, Iain; Pickering, David A. et al. (February 1, 2011). "A diverse Pleistocene marsupial trackway assemblage from the Victorian Volcanic Plains, Australia". Quarternary Science Reviews 30 (5-6): 591–610. doi:10.1016/j.quascirev.2010.11.021. Retrieved 18 October 2014.
- Camens, Aaron Bruce; Carey, Stephen Paul (January 1, 2013). "Contemporaneous Trace and Body Fossils from a Late Pleistocene Lakebed in Victoria, Australia Allow Assessment of Bias in the Fossil Record". PLoS One 8 (1): e52957. doi:10.1371/journal.pone.0052957. Retrieved 18 October 2014.
- "The Extinction Enigma". PBS. May 1, 2007.
- Anatomy of the Marsupial Lion (Interactive feature from Nova)
- BBC news article about Wroe's comparative study of bite force amongst mammalian predators
- Fact sheet from Australian Museum Online[dead link]
- Move Over Sabre-Tooth Tiger by Stephen Wroe from Australian Museum Online.[dead link]
- The Extinction Enigma (Interview from Nova)