|Diprotodon skeleton cast, MNHN, Paris|
Diprotodon is an extinct genus of large marsupials native to Australia from the Pleistocene epoch. It is considered one of Australia's core species of "megafauna", which ranged throughout the continent during the Pleistocene. The genus is currently considered monotypic, containing only Diprotodon optatum, the largest known marsupial to have ever existed. The word diprotodon is constructed from the Ancient Greek words for 'two forward teeth'. Diprotodon existed from about 1.6 million years ago until extinction some 44,000 years ago.
Diprotodon species fossils have been found in sites across mainland Australia,[a] including complete skulls, skeletons, and foot impressions.[better source needed] The largest specimens were hippopotamus-sized: about 3 m (9.8 ft) from nose to tail, standing 2 m (6.6 ft) tall at the shoulder and weighing about 2,790 kg (6,150 lb).[b] Diprotodons may have been depicted on Aboriginal rock art images in Quinkan traditional country (Queensland, Australia). Diprotodon became extinct sometime after 44,000 years ago, after the initial settlement of the continent; the role of human and climatic factors in its extinction are uncertain and contested.
Diprotodon is a member of the extinct family Diprotodontidae. The closest surviving relatives of Diprotodon are the wombats and the koala, and so Diprotodon have sometimes been referred to as "giant wombats" in the popular press. Diprotodonts are suggested to have inspired legends of the bunyip, as some Aboriginal tribes identify Diprotodon bones as those of "bunyips".[c]
In 1830, bushman George Ranken found a diverse fossil assemblage while exploring Wellington Caves, New South Wales, and was the first major site of extinct Australian megafauna. They were excavated when Ranken returned later in a formal expedition headed by Major Thomas Mitchell.
The majority of fossil finds are of demographic groups indicative of diprotodonts dying in drought conditions. For example, hundreds of individuals were found in Lake Callabonna with well-preserved lower bodies, but crushed and distorted heads. Several family groups are thought to have sunk in mud while crossing the drying lake bed. Other finds consist of age groupings of young or old animals, which are first to die during a drought.
At the time these massive fossils were discovered, there were no serious scientists in Australia, so the massive fossils were not formally described until Mitchell, while in England publishing his journal, brought them to his former colleague Sir Richard Owen in 1837. Before Owen, it was generally guessed the fossil assemblage represents rhinos, elephants, hippos, or dugongs. In 1838, while studying a piece of a right mandible with an incisor, Owen compared the tooth to that of a wombat or a hippo, and designated it as a new genus in a letter to Mitchell, as Diprotodon. Mitchell published the correspondence in his journal. He formally described Diprotodon in volume 2 without mentioning a species, but in volume 1, he listed the name Diprotodon optatum, making that the type species. Diprotodon means "two protruding front teeth". In 1844, Owen replaced the name D. optatum with "D. australis". Owen only used optatum the one time, and the acceptance of its apparent replacement "australis" has varied widely.
In 1843, Mitchell was sent more Diprotodon fossils from the recently settled the Darling Downs, and relayed them to Owen. Owen believed Diprotodon was an elephant related to or synonymous with Mastodon or Deinotherium, having interpreted the incisors as tusks, as well as comparing the flattening (anteroposterior compression) of the femur to the condition in elephants and rhinos, and the raised ridges of the molar to the grinding surfaces of elephant teeth. Later that year, he formally synonymised Diprotodon with Deinotherium as "Dinotherium Australe", which he recanted in 1844 after German naturalist Ludwig Leichhardt pointed out that the incisors clearly belong to a marsupial. Owen, nonetheless, still classified the molars from Wellington as "M. australis", and described Diprotodon as elephantine. In 1847, a nearly complete skull and skeleton was recovered from the Darling Downs, which confirmed this characterisation. Leichhardt suggested the animal still may be alive judging by the quality of preservation, but as the European land exploration of Australia progressed, he became certain it was extinct.
Several other species were subsequently erected because adult Diprotodon specimens come in two distinct size ranges. This was not originally ascribed to sexual dimorphism because males and females among modern wombat and koala species are skeletally indistinguishable, so it was assumed the same would have been true for extinct relatives, including Diprotodon. These other species are:
- "D. annextans", erected in 1861 by Irish palaeontologist Frederick McCoy based on some teeth and a partial mandible near Colac, Victoria; the name may be a typo of annectens which means linking or joining, because the species combines traits from Diprotodon and Nototherium;
- "D. minor", erected in 1862 by Thomas Huxley based on a partial palate; in 1999, Australian palaeontologist Peter Murray suggested classifying large specimens as D. optatum and smaller ones as "D. minor";
- "D. longiceps", erected in 1865 by McCoy as a replacement for "D. annextans";
- "D. bennettii", erected in 1873 by German naturalist Gerard Krefft based on a nearly complete mandible collected by naturalists George Bennet and Georgina King near Gowrie, New South Wales;
- and "D. loderi", erected in 1873 by Krefft based on a partial palate collected by Andrew Loder near Murrurundi, New South Wales.
In 2008, Australian palaeontologist Gilbert Price opted to recognise only one species, D. optatum.
Diprotodon is ultimately thought to have evolved from Euryzygoma dunensis, a smaller diprotodontid known from the Pliocene of eastern Australia, and a smaller form of Diprotodon, labelled D. ?optatum, intermediate in size between the two taxa, is known from the Early Pleistocene (1.77–0.78 Ma) in Nelson Bay near Portland, Victoria.
Diprotodon superficially resembled a giant wombat. The length of the skull of adult Diprotodon ranged between 65 cm and 1 m. In side view, the cranial profile is elongate and low. The zygomatic arch and flare laterally. Its feet turned inwards like a wombat's, giving it a pigeon-toed stance. It had strong claws on the front feet and its pouch opening faced backwards.
John Walter Gregory collected stories of mystical creatures in Aboriginal myths and legends and considered the possible connections between them and extinct species. He reported a story of a "big, heavy land animal, with a single horn on its forehead" as a possible reference to Diprotodon; the presence of a horn on the rostrum of the species is not scientifically acknowledged.
The skull of Diprotodon has large endocranial sinus cavities, which separate the relatively small cranial vault from the outer part of the skull. These significantly lighten the skull while providing large areas for muscle attachment and reduce load stress.
Recent research compared the variation between all of the described Diprotodon species with the variation in one of Australia's largest living marsupials, the eastern grey kangaroo, and found the range was comparable, with a near continent-wide distribution. This left only two possible Diprotodon species differing only in size with the smaller being around half the size of the larger. According to Gause's "competitive exclusion principle", no two species with identical ecological requirements can coexist in a stable environment. However, both the small and large diprotodonts coexisted throughout the Pleistocene and the size difference is similar to other sexually dimorphic living marsupials. Further evidence is the battle damage common in competing males found on the larger specimens, but absent from the smaller. Dental morphology also supports sexual dimorphism, with highly sexually dimorphic marsupials, such as the grey kangaroo, having different tooth sizes between males and females, but both sexes having the same dental morphology. An identical dental morphology occurs in the large and small Diprotodon. The taxonomic implication is that Owen's original Diprotodon optatum is the only valid species.
A single sexually dimorphic species allows behavioural interpretations. All sexually dimorphic species of over 5 kg (11 lb) exhibit a polygynous breeding strategy. A modern example of this is the gender segregation of elephants, where females and the young form family groups, while lone males fight for the right to mate with all the females of the group. This behaviour is consistent with fossil finds where adult/juvenile fossil assemblages usually contain only female adult remains.
Cyclic variations in the strontium isotope ratios within the tooth enamel of a 300,000-year-old fossil imply that a population of Diprotodon undertook regular, seasonal migrations across the Darling Downs, making it the only known extinct or extant metatherian known to migrate annually. The carbon-13 enamel content was found to have little variation, suggesting a relatively consistent diet through the course of a year containing a mix of both C3 and C4 plants. A finite element method analysis of the skull estimated it had a bite force of around 4,500 Newtons at the first molar to over 11,000 N at the fourth molar, values which were described as "exceptionally high", suggesting that Diprotodon was capable of processing tough, fibrous food.
Some modern researchers, including Richard Roberts and Tim Flannery, argue that diprotodonts, along with a wide range of other Australian megafauna, became extinct shortly after humans arrived in Australia about 50,000 years ago. Others, including Steve Wroe, note that records in the Australian Pleistocene are rare, and there is not enough data to definitively determine the time of extinction of many of the species, with many of the species having no confirmed record within the last 100,000 years. They suggest that many of the extinctions had been staggered over the course of the late Middle Pleistocene and early Late Pleistocene, prior to human arrival, due to climatic stress. Diprotodon is one of several species with confirmed dates post-dating human arrival on the continent, with the latest high-reliability date being around 44,000 years Before Present.
Some earlier researchers, including Richard Wright, argued on the contrary that diprotodont remains from several sites, such as Tambar Springs and Trinkey and Lime Springs, suggest that Diprotodon survived much longer, into the Holocene. Other more recent researchers, including Lesley Head and Judith Field, favour an extinction date of 28,000–30,000 years ago, which would mean that humans coexisted with Diprotodon for some 20,000 years. However, opponents of "late extinction" theories have interpreted such late dates based on indirect dating methods as artifacts resulting from redeposition of skeletal material into more recent strata, and recent direct dating results obtained with new technologies have tended to confirm this interpretation.
Three theories have been advanced to explain the mass extinction.
Australia, as with many other areas of the globe, underwent significant ecological change from predominantly forests to open landscapes during the late Neogene, in correlation with increased cooling. There has been considerable aridification of the Australian interior since the Late Miocene. The recent ice ages produced no significant glaciation in mainland Australia, but long periods of cold and increased aridification. These climatic changes have been suggested as a contributing factor in extinction.
The overkill theory is that human hunters killed and ate the diprotodonts, causing their extinction. The extinctions appear to have coincided with the arrival of humans on the continent. Similar hunting-out happened with the megafauna of New Zealand, Madagascar, and many other islands around the world, such as New Caledonia and the Greater Antilles. Critics of this theory regard it as simplistic, arguing that (unlike New Zealand and America), little direct evidence of hunting has been found, and the dates on which the theory rests are too uncertain to be reliable. Linear striated cuts on a Diprotodon tooth from south-eastern Australia initially suggested to have been etched by people are now thought to be bite marks from a spotted‐tailed quoll. A partial juvenile radius of Diprotodon optatum over 47,000 years old was found at the Warratyi rock shelter in South Australia north of Adelaide. Due to the shelter's location on a steep escarpment and the lack of bite marks on the bone, it is thought that humans transported the bone to the site. At present, this represents the only known interaction between humans and Diprotodon.
An examination of swamp sediment cores spanning the last 130,000 years from Lynch's Crater in Queensland suggests that hunting may have been the primary cause of the extinction. Analysis of Sporormiella fungal spores (which are found in the dung of herbivores and are used as a proxy for their abundance) in the cores shows that the Spororniella records in that region virtually disappeared about 41,000 years ago, at a time when climate changes were minimal; the change was accompanied by an increase in charcoal, and was followed by a transition from rainforest to fire-tolerant sclerophyll vegetation. The high-resolution chronology of the changes indicates that fire increased about a century after the disappearance of browsing megafauna, probably due to accumulation of fuel. Grass increased over the next several centuries; sclerophyll vegetation increased following a lag of another century, and a sclerophyll forest developed about a thousand years later. Earlier increases in sclerophyll vegetation during shifts to cooler, drier conditions about 120,000 and 75,000 years ago did not have any obvious impact on megafaunal abundance. A study on Sporormiella in cores going back 135,000 years at the Caledonia Fen wetland in Alpine National Park, found that Sporomiella levels rose after the end of the last interglacial, followed by a sharp drop around 76–60 kya, associated with an increased proportion of wetland plants and further grassland and herbfield; a second sharp drop was observed around 52–45 kya, suggested to represent the megafaunal extinction interval. Charcoal was found throughout the core, suggesting that the arrival of humans and the extinction of megafauna did not change the fire regime at this locality. However, the use of Sporormiella as a megafaunal proxy has been criticised, noting that Sporormiella is found sporadically in the dung of various herbivores, including extant emus and kangaroos, not just megafauna, its presence depends on a variety of factors, often unrelated to megafaunal abundance, and that in Cuddie Springs, a well known megafaunal site, the densities of Sporormiella were consistently low.
Human land management
The third theory says that humans indirectly caused the extinction of diprotodonts by destroying the ecosystem on which they depended. In particular, early Aboriginal people are thought to have been fire-stick farmers using fire regularly and persistently to drive game, open up dense thickets of vegetation, and create fresh green regrowth for both humans and game animals to eat. Evidence for the fire hypothesis is the sudden increase in widespread ash deposits at the time that people arrived in Australia, as well as land-management and hunting practices of modern Aboriginal people as recorded by the earliest European settlers. Evidence against the hypothesis is the fact that humans appear to have eliminated the megafauna of Tasmania without using fire to modify the environment there.
- Including Kangaroo Island
- The estimated mean body mass for the 17 adult specimens studied, calculated from the minimum midshaft circumferences of the femur and humerus, is 2,786 kg, although the size of these mammals is believed to be larger because the cartilage is not being accounted for when they fossilize. The 95% confidence interval for this value ranges from 2,272 kg to 3,417 kg.
- "As far back as 1924, Dr. C.W. Anderson of the Australian Museum had suggested that stories of the bunyip could derive from aboriginal legends of the extinct diprotodonts – a view repeated much more recently in Kadimakara (1985) by Australian zoologists Drs. Tim Flannery and Michael Archer, who nominated the palorchestids as plausible candidates."
— K.P.N. Shuker (1995)
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|Wikispecies has information related to Diprotodon.|
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|Wikimedia Commons has media related to Diprotodon.|
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