Thylacine

The thylacine (/ˈθaɪləsiːn/ THY-lə-seen,[14] or /ˈθaɪləsaɪn/ THY-lə-syne,[15] also /ˈθaɪləsɪn/)[16] (Thylacinus cynocephalus) is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea.[17] The last known live animal was captured in 1930 in Tasmania. It is commonly known as the Tasmanian tiger (because of its striped lower back) or the Tasmanian wolf (because of its canid-like characteristics). Various Aboriginal Tasmanian names have been recorded, such as coorinna, kanunnah, cab-berr-one-nen-er, loarinna, laoonana, can-nen-ner and lagunta,[18][19] while kaparunina is used in Palawa kani.[20][21]

Thylacine[1]
Temporal range: Pleistocene–Holocene,
A female thylacine and her juvenile offspring in the National Zoo in Washington, D.C., c. 1903[3]

Extinct (1936)  (IUCN 3.1)[4]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Marsupialia
Order: Dasyuromorphia
Family: †Thylacinidae
Genus: †Thylacinus
Species:
†T. cynocephalus
Binomial name
†Thylacinus cynocephalus
(Harris, 1808)[5]
Historic thylacine range in Tasmania (in green)[6]
Synonyms
List
  • Didelphis cynocephala Harris, 1808[5]
  • Dasyurus cynocephalus Geoffroy, 1810[7]
  • Thylacinus harrisii Temminck, 1824[8]
  • Dasyurus lucocephalus Grant, 1831[9]
  • Thylacinus striatus Warlow, 1833[10]
  • Thylacinus communis Anon., 1859[11]
  • Thylacinus breviceps Krefft, 1868[12]
  • Thylacinus rostralis De Vis, 1893[13]

The thylacine was relatively shy and nocturnal, with the general appearance of a medium-to-large-size canid, except for its stiff tail and abdominal pouch similar to that of a kangaroo. Because of convergent evolution, it displayed an anatomy and adaptations similar to the tiger (Panthera tigris) and wolf (Canis lupus) of the Northern Hemisphere, such as dark transverse stripes that radiated from the top of its back, and a skull shape extremely similar to those of canids, despite being unrelated. The thylacine was a formidable apex predator,[6] though exactly how large its prey animals were is disputed. Its closest living relatives are the other members of Dasyuromorphia, including the Tasmanian devil and quolls. The thylacine was one of only two marsupials known to have a pouch in both sexes: the other (still extant) species is the water opossum from Central and South America. The pouch of the male thylacine served as a protective sheath, covering the external reproductive organs.

The thylacine had become locally extinct on both New Guinea and the Australian mainland before British settlement of the continent, but its last stronghold was on the island of Tasmania, along with several other endemic species, including the Tasmanian devil. Intensive hunting encouraged by bounties is generally blamed for its extinction, but other contributing factors may have been disease, the introduction of and competition with dingoes, and human encroachment into its habitat.

Taxonomic and evolutionary history
Tasmanian devil and thylacine, both labelled as members of Didelphis, from Harris' 1808 description. This is the earliest known non-indigenous illustration of a thylacine.

Numerous examples of thylacine engravings and rock art have been found, dating back to at least 1000 BC.[22] Petroglyph images of the thylacine can be found at the Dampier Rock Art Precinct, on the Burrup Peninsula in Western Australia.

By the time the first European explorers arrived, the animal was already extinct in mainland Australia and New Guinea, and rare in Tasmania. Europeans may have encountered it in Tasmania as far back as 1642, when Abel Tasman first arrived in Tasmania. His shore party reported seeing the footprints of "wild beasts having claws like a Tyger".[23] Marc-Joseph Marion du Fresne, arriving with the Mascarin in 1772, reported seeing a "tiger cat".[24] Positive identification of the thylacine as the animal encountered cannot be made from this report, since the tiger quoll (Dasyurus maculatus) is similarly described.

The first definitive encounter was by French explorers on 13 May 1792, as noted by the naturalist Jacques Labillardière, in his journal from the expedition led by d'Entrecasteaux. In 1805, William Paterson, the Lieutenant Governor of Tasmania, sent a detailed description for publication in the Sydney Gazette.[25] He also sent a description of the thylacine in a letter to Joseph Banks, dated 30 March 1805.[26]

The first detailed scientific description was made by Tasmania's Deputy Surveyor-General, George Harris, in 1808, five years after first European settlement of the island.[5][27][28] Harris originally placed the thylacine in the genus Didelphis, which had been created by Linnaeus for the American opossums, describing it as Didelphis cynocephala, the "dog-headed opossum". Recognition that the Australian marsupials were fundamentally different from the known mammal genera led to the establishment of the modern classification scheme, and in 1796, Geoffroy Saint-Hilaire created the genus Dasyurus, where he placed the thylacine in 1810. To resolve the mixture of Greek and Latin nomenclature, the species name was altered to cynocephalus. In 1824, it was separated out into its own genus, Thylacinus, by Temminck.[29] The common name derives directly from the genus name, originally from the Greek θύλακος (thýlakos), meaning "pouch" or "sack".[30][a]

Evolution
The skulls of the thylacine (left) and the grey wolf (Canis lupus) are quite similar, although the species are not related. Studies show that the skull shape of the red fox (Vulpes vulpes), is even closer to that of the thylacine.[31]

The modern thylacine probably appeared about 2 million years ago, during the Early Pleistocene. Specimens from the Pliocene-aged Chinchilla Fauna, described as Thylacinus rostralis by Charles De Vis in 1894, have in the past been suggested to represent Thylacinus cynocephalus, but have been shown to either have been curatorial errors, or ambiguous in their specific attribution.[32][33] The family Thylacinidae includes at least 12 species in eight genera,[34] and appears around the late Oligocene with the small, plesiomorphic Badjcinus turnbulli.[35] Early thylacinids were quoll-sized, well under 10 kg (22 lb), and probably ate insects and small reptiles and mammals, although signs of an increasingly-carnivorous diet can be seen as early as the early Miocene in Wabulacinus.[34] Members of the genus Thylacinus are notable for a dramatic increase in both the expression of carnivorous dental traits and in size, with the largest species, Thylacinus potens and Thylacinus megiriani both approaching the size of a wolf.[34] In Late Pleistocene and early Holocene times, the modern thylacine was widespread (although never numerous) throughout Australia and New Guinea.[36]

A classic example of convergent evolution, the thylacine showed many similarities to the members of the dog family, Canidae, of the Northern Hemisphere: sharp teeth, powerful jaws, raised heels, and the same general body form. Since the thylacine filled the same ecological niche in Australia and New Guinea as canids did elsewhere, it developed many of the same features. Despite this, as a marsupial, it is unrelated to any of the Northern Hemisphere placental mammal predators.[37]

They are easy to tell from a true dog because of the stripes on the back but the skeleton is harder to distinguish. Zoology students at Oxford had to identify 100 zoological specimens as part of the final exam. Word soon got around that, if ever a 'dog' skull was given, it was safe to identify it as Thylacinus on the grounds that anything as obvious as a dog skull had to be a catch. Then one year the examiners, to their credit, double bluffed and put in a real dog skull. The easiest way to tell the difference is by the two prominent holes in the palate bone, which are characteristic of marsupials generally.[38]

— Richard Dawkins, The Ancestor's Tale

The thylacine is a basal member of the Dasyuromorphia, along with numbats, dunnarts, wambengers, and quolls. The cladogram follows:[39]

Dasyuromorphia
 

Thylacinus (thylacines)

 
 
 

Myrmecobius (numbat)

 
 
 

Sminthopsis (dunnarts)

 
 
 

Phascogale (wambengers)

 
 

Dasyurus (quolls)

 
 
 
 
 

Description

The only recorded species of Thylacinus, a genus that superficially resembles the dogs and foxes of the family Canidae, the animal was a predatory marsupial that existed on mainland Australia during the Holocene epoch and observed by Europeans on the island of Tasmania; the species is known as the Tasmanian tiger for the striped markings of the pelage. Descriptions of the thylacine come from preserved specimens, fossil records, skins and skeletal remains, and black and white photographs and film of the animal both in captivity and from the field. The thylacine resembled a large, short-haired dog with a stiff tail which smoothly extended from the body in a way similar to that of a kangaroo.[37] The mature thylacine ranged from 100 to 130 cm (39 to 51 in) long, plus a tail of around 50 to 65 cm (20 to 26 in).[40] Adults stood about 60 cm (24 in) at the shoulder and on average weighed 12 to 22 kg (26 to 49 lb), though they could range anywhere from 8 to 30 kg (18 to 66 lb).[41] There was slight sexual dimorphism with the males being larger than females on average.[42] Males weighed in at around 19.7 kilograms (43 lb), and females weighed in at around 13.7 kilograms (30 lb).[41][43] The skull is noted to be highly convergent on those of canids, most closely remembling that of the red fox.[44]

Thylacines, uniquely for marsupials, had largely cartilaginous epipubic bones with a highly reduced osseous element.[45][46] This has been once considered a synapomorphy with sparassodonts,[47] though it is now thought that both groups reduced their epipubics independently. Its yellow-brown coat featured 15 to 20 distinctive dark stripes across its back, rump and the base of its tail,[48] which earned the animal the nickname "tiger". The stripes were more pronounced in younger specimens, fading as the animal got older.[48] One of the stripes extended down the outside of the rear thigh. Its body hair was dense and soft, up to 15 mm (0.6 in) in length. Colouration varied from light fawn to a dark brown; the belly was cream-coloured.[49]

Its rounded, erect ears were about 8 cm (3.1 in) long and covered with short fur.[50] The early scientific studies suggested it possessed an acute sense of smell which enabled it to track prey,[51] but analysis of its brain structure revealed that its olfactory bulbs were not well developed. It is likely to have relied on sight and sound when hunting instead.[48]

The thylacine was able to open its jaws to an unusual extent: up to 80 degrees.[52] This capability can be seen in part in David Fleay's short black-and-white film sequence of a captive thylacine from 1933. The jaws were muscular, and had 46 teeth, but studies show the thylacine jaw was too weak to kill sheep.[50][53][54] The tail vertebrae were fused to a degree, with resulting restriction of full tail movement. Fusion may have occurred as the animal reached full maturity. The tail tapered towards the tip. In juveniles, the tip of the tail had a ridge.[55] The female thylacine had a pouch with four teats, but unlike many other marsupials, the pouch opened to the rear of its body. Males had a scrotal pouch, unique amongst the Australian marsupials,[56] into which they could withdraw their scrotal sac for protection.[48]

The thylacine's footprint is easy to distinguish from those of native and introduced species.

Thylacine footprints could be distinguished from other native or introduced animals; unlike foxes, cats, dogs, wombats, or Tasmanian devils, thylacines had a very large rear pad and four obvious front pads, arranged in almost a straight line.[51] The hindfeet were similar to the forefeet but had four digits rather than five. Their claws were non-retractable.[48] More detail can be seen in a cast taken from a freshly dead thylacine. The cast shows the plantar pad in more detail and shows that the plantar pad is tri-lobal in that it exhibits three distinctive lobes. It is a single plantar pad divided by three deep grooves. The distinctive plantar pad shape along with the asymmetrical nature of the foot makes it quite different from animals such as dogs or foxes. This cast dates back to the early 1930s and is part of the Museum of Victoria's thylacine collection.[57]

The thylacine was noted as having a stiff and somewhat awkward gait, making it unable to run at high speed. It could also perform a bipedal hop, in a fashion similar to a kangaroo—demonstrated at various times by captive specimens.[48] Guiler speculates that this was used as an accelerated form of motion when the animal became alarmed.[49] The animal was also able to balance on its hind legs and stand upright for brief periods.[58]

Observers of the animal in the wild and in captivity noted that it would growl and hiss when agitated, often accompanied by a threat-yawn. During hunting, it would emit a series of rapidly repeated guttural cough-like barks (described as "yip-yap", "cay-yip" or "hop-hop-hop"), probably for communication between the family pack members. It also had a long whining cry, probably for identification at distance, and a low snuffling noise used for communication between family members.[59] Some observers described it as having a strong and distinctive smell, others described a faint, clean, animal odour, and some no odour at all. It is possible that the thylacine, like its relative, the Tasmanian devil, gave off an odour when agitated.[60]

Distribution and habitat

The thylacine most likely preferred the dry eucalyptus forests, wetlands, and grasslands of mainland Australia.[51] Indigenous Australian rock paintings indicate that the thylacine lived throughout mainland Australia and New Guinea. Proof of the animal's existence in mainland Australia came from a desiccated carcass that was discovered in a cave in the Nullarbor Plain in Western Australia in 1990; carbon dating revealed it to be around 3,300 years old.[61] Recently examined fossilised footprints also suggest historical distribution of the species on Kangaroo Island.[62] The northernmost record of the species is from the Kiowa rock shelter in Chimbu Province in the highlands of Papua New Guinea, dating to the Early Holocene, ~10,000-8,500 years Before Present.[63]

In Tasmania it preferred the woodlands of the midlands and coastal heath, which eventually became the primary focus of British settlers seeking grazing land for their livestock.[64] The striped pattern may have provided camouflage in woodland conditions,[48] but it may have also served for identification purposes.[65] The animal had a typical home range of between 40 and 80 km2 (15 and 31 sq mi).[49] It appears to have kept to its home range without being territorial; groups too large to be a family unit were sometimes observed together.[66]

Ecology and behaviour
One of only two known photos of a thylacine with a distended pouch, bearing young. Hobart Zoo, 1928
Thylacine family at Beaumaris Zoo in Hobart, 1909
Thylacine family at Beaumaris Zoo in Hobart, 1910

Little is known about the behaviour of the thylacine. A few observations were made of the animal in captivity, but only limited, anecdotal evidence exists of the animal's behaviour in the wild. Most observations were made during the day, whereas the thylacine was naturally nocturnal. Those observations, made in the twentieth century, may have been atypical, as they were of a species already under the stresses that would soon lead to its extinction. Some behavioural characteristics have been extrapolated from the behaviour of its close relative, the Tasmanian devil.

The thylacine was a nocturnal and crepuscular hunter, spending the daylight hours in small caves or hollow tree trunks in a nest of twigs, bark, or fern fronds. It tended to retreat to the hills and forest for shelter during the day. and hunted in the open heath at night. Early observers noted that the animal was typically shy and secretive, with awareness of the presence of humans and generally avoiding contact, though it occasionally showed inquisitive traits.[67] At the time, much stigma existed in regard to its "fierce" nature; this is likely to be due to its perceived threat to agriculture.[68]

Reproduction

There is evidence for at least some year-round breeding (cull records show joeys discovered in the pouch at all times of the year), although the peak breeding season was in winter and spring.[48] They would produce up to four joeys per litter (typically two or three), carrying the young in a pouch for up to three months and protecting them until they were at least half adult size. Early pouch young were hairless and blind, but they had their eyes open and were fully furred by the time they left the pouch.[69] The young also have their own pouches but are not visible until they are 9.5 weeks old.[48] After leaving the pouch, and until they were developed enough to assist, the juveniles would remain in the lair while their mother hunted.[70] Thylacines only once bred successfully in captivity, in Melbourne Zoo in 1899.[71] Their life expectancy in the wild is estimated to have been 5 to 7 years, although captive specimens survived up to 9 years.[51]

In 2018, Newton et al. collected and CT-scanned all known preserved thylacine pouch young specimens to digitally reconstruct its development throughout its entire window of growth in the mother's pouch. This study revealed new information on the biology of the thylacine, including the growth of its limbs and when it developed its 'dog-like' appearance. It was found that two of the thylacine young in the Tasmanian Museum and Art Gallery (TMAG) were misidentified and of another species, reducing the number of known pouch young specimens to 11 worldwide.[72] One of four specimens kept at Museum Victoria has been serially sectioned, allowing an in-depth investigation of its internal tissues and providing some insights into thylacine pouch young development, biology, immunology and ecology.[73]

Feeding and diet
1887 illustration of an emu being chased by two thylacines

The thylacine was exclusively carnivorous. In captivity, thylacines had a clear preference for birds (particularly chickens). In the wild, large ground-dwelling birds (such as Tasmanian nativehens) may have been their primary prey, since they are documented to have hunted a wide range of them, and its comparatively moderate bite force was more suited to hollow avian bones. During its peak occupation of the mainland, such prey would have been bountiful, and studies of their Pleistocene habitat points to a more suitable diet consisting of a range of megapodes (such as the Giant malleefowl)[74] ratites (such as the emu), and possibly dromornithids (most of which became extinct prior to European settlement). At the time of European settlement, the Tasmanian emu, a subspecies believed to be smaller than mainland emus, was common and widespread and Thylacines were known to prey on them and share the same habitat.[75] Many early depictions of them hunting included emu. The large, flightless bird was hunted to extinction by humans within 30 years of European settlement. The extinction correlates with a rapid decline in thylacine numbers.[76][77][78] Cassowary species of northern Australia and New Guinea coexisted with the Thylacine, but had developed strong defenses against predators;[79] the emu on the other hand was more vulnerable to the Thylacine's adaptions including endurance hunting and bipedal hop.[78] Dingoes, feral dogs, and red foxes have all been noted to hunt the emu on the mainland[80][81] and killings of emus by dogs were noted in Tasmania.[82] European settlers believed the thylacine to prey upon farmers' sheep and poultry.[83][84] Throughout the 20th century, the thylacine was often characterised as primarily a blood drinker; according to Robert Paddle, the story's popularity seems to have originated from a single second-hand account heard by Geoffrey Smith (1881–1916)[85][86] in a shepherd's hut.[87]

Analysis of the skeleton suggests that, when hunting, the thylacine relied on stamina rather than speed in the chase.

There is some controversy over the preferred prey size of the thylacine. A 2011 study by the University of New South Wales using advanced computer modelling indicated that the thylacine had surprisingly feeble jaws. Animals usually take prey close to their own body size, but an adult thylacine of around 30 kilograms (66 lb) was found to be incapable of handling prey much larger than 5 kilograms (11 lb). Thus, some researchers believe thylacines only ate small animals such as bandicoots and possums, putting them into direct competition with the Tasmanian devil and the tiger quoll. Another study in 2020 produced similar results, after estimating the average thylacine weight as about 17 kilograms (37 lb) rather than 30 kilograms (66 lb), suggesting that the animal did indeed hunt much smaller prey.[43]

However, an earlier study showed that the thylacine had a bite force quotient of 166, similar to that of most quolls; in modern mammalian predators, such a high bite force is almost always associated with predators which routinely take prey as large, or larger than, themselves.[88] If the thylacine was indeed specialised for small prey, this specialisation likely made it susceptible to small disturbances to the ecosystem.[89]

Analysis of the skeletal frame and observations of the thylacine in captivity suggest the species were pursuit predators, singling out a prey item and pursuing them until the prey was exhausted. However, trappers reported it as an ambush predator.[48] The animal may have hunted in small family groups, with the main group herding prey in the general direction of an individual waiting in ambush.[27] Although the living grey wolf is widely seen as the thylacine's counterpart, the thylacine may have been more of an ambush predator as opposed to a pursuit predator. In fact, the predatory behaviour of the thylacine was probably closer to ambushing felids than to large pursuit canids. Its stomach was muscular, and could distend to allow the animal to eat large amounts of food at one time, probably an adaptation to compensate for long periods when hunting was unsuccessful and food scarce.[48]

In captivity, thylacines were fed a wide variety of foods, including dead rabbits and wallabies as well as beef, mutton, horse, and occasionally poultry.[90] There is a report of a captive thylacine which refused to eat dead wallaby flesh or to kill and eat a live wallaby offered to it, but "ultimately it was persuaded to eat by having the smell of blood from a freshly killed wallaby put before its nose."[91]

In 2017, Berns and Ashwell published comparative cortical maps of thylacine and Tasmanian devil brains, showing that the thylacine had a larger, more modularised basal ganglion. The authors associated these differences with the thylacine's predatory lifestyle.[92] The same year, White, Mitchell and Austin published a large-scale analysis of thylacine mitochondrial genomes, showing that they had split into Eastern and Western populations on the mainland prior to the Last Glacial Maximum and had low genetic diversity by the time of European arrival.[93]

Relationship with humans
Thylacine rock art at Ubirr

Captivity

By the beginning of the 20th century, the increasing rarity of thylacines led to increased demand for captive specimens by zoos around the world.[94] Despite the export of breeding pairs, attempts at having thylacines in captivity were unsuccessful, and the last thylacine outside Australia died at the London Zoo in 1931.[95]

Extinction in the Australian mainland
Killed thylacine, 1869

Australia lost more than 90% of its megafauna by around 40,000 years ago, with the notable exceptions of several kangaroo species and the thylacine.[96] A 2010 paper examining this issue showed that humans were likely to be one of the major factors in the extinction of many species in Australia although the authors of the research warned that one-factor explanations might be over-simplistic.[96] The thylacine itself likely neared extinction throughout most of its range in mainland Australia by about 2,000 years ago.[4]

However, reliable accounts of thylacine survival in South Australia (though confined to the "thinly settled districts" and Flinders Ranges) and New South Wales (Blue Mountains) exist from as late as the 1830s, from both indigenous and European sources.[97]

A study proposes that the arrival of the dingoes may have led to the extinction of the Tasmanian devil, the thylacine, and the Tasmanian native hen in mainland Australia because the dingo might have competed with the thylacine and devil in preying on the native hen. The study also proposes that an increase in the human population that gathered pace around 4,000 years ago may have led to this.[98]

A counter-argument is that the two species were not in direct competition with one another because the dingo primarily hunts during the day, whereas it is thought that the thylacine hunted mostly at night. Nonetheless, recent morphological examinations of dingo and thylacine skulls show that although the dingo had a weaker bite, its skull could resist greater stresses, allowing it to pull down larger prey than the thylacine. The thylacine was less versatile in its diet than the omnivorous dingo.[99][100] Their ranges appear to have overlapped because thylacine subfossil remains have been discovered near those of dingoes. The adoption of the dingo as a hunting companion by the indigenous peoples would have put the thylacine under increased pressure.[98]

A 2013 study suggested that, while dingoes were a contributing factor to the thylacine's demise on the mainland around 3,000 years ago, larger factors were the intense human population growth and technological advances and the abrupt change in the climate during the period.[101][102]

Extinction in Tasmania

Although the thylacine was extinct on mainland Australia, it survived into the 1930s on the island state of Tasmania. At the time of the first European settlement, the heaviest distributions were in the northeast, northwest and north-midland regions of the state.[64] They were rarely sighted during this time but slowly began to be credited with numerous attacks on sheep. This led to the establishment of bounty schemes in an attempt to control their numbers. The Van Diemen's Land Company introduced bounties on the thylacine from as early as 1830, and between 1888 and 1909 the Tasmanian government paid £1 per head for dead adult thylacines and ten shillings for pups. In all, they paid out 2,184 bounties, but it is thought that many more thylacines were killed than were claimed for. Its extinction is popularly attributed to these relentless efforts by farmers and bounty hunters.[51][103][104]

Tasmanian tiger skeletons on display at Tasmanian Museum and Art Gallery Hobart

However, it is likely that multiple factors led to its decline and eventual extinction, including competition with wild dogs introduced by European settlers,[105] erosion of its habitat, the concurrent extinction of prey species, and a distemper-like disease that affected many captive specimens at the time.[49][106] A study from 2012 also found that were it not for an epidemiological influence, the extinction of thylacine would have been at best prevented, at worst postponed. "The chance of saving the species, through changing public opinion, and the re-establishment of captive breeding, could have been possible. But the marsupi-carnivore disease, with its dramatic effect on individual thylacine longevity and juvenile mortality, came far too soon, and spread far too quickly."[107]

Whatever the reason, the animal had become extremely rare in the wild by the late 1920s. Despite the fact that the thylacine was believed by many to be responsible for attacks on sheep, in 1928 the Tasmanian Advisory Committee for Native Fauna recommended a reserve similar to the Savage River National Park to protect any remaining thylacines, with potential sites of suitable habitat including the Arthur-Pieman area of western Tasmania.[108]

The last known thylacine to be killed in the wild was shot in 1930 by Wilf Batty, a farmer from Mawbanna in the state's northwest. The animal, believed to have been a male, had been seen around Batty's house for several weeks.[109][110]

Work in 2012 examined the relationship of the genetic diversity of the thylacines before their extinction. The results indicated that the last of the thylacines in Tasmania had limited genetic diversity due to their complete geographic isolation from mainland Australia.[111] Further investigations in 2017 showed evidence that this decline in genetic diversity started long before the arrival of humans in Australia, possibly starting as early as 70–120 thousand years ago.[44]

  • This 1921 photo by Henry Burrell of a thylacine with a chicken was widely distributed and may have helped secure the animal's reputation as a poultry thief. In fact the image is cropped to hide the fenced run and housing, and analysis by one researcher has concluded that this thylacine is a mounted specimen, posed for the camera. The photograph may even have involved photo manipulation.[upper-alpha 1]
  • Wilf Batty with the last thylacine that was killed in the wild.
  • Thylacine (Tasmanian tiger) trap, intended for Mount Morriston, 1823, by Thomas Scott

Benjamin and searches
Benjamin was an endling, and lived as the last known Tasmanian tiger, photographed at Hobart Zoo in 1933.
The last known thylacine photographed at Beaumaris Zoo in 1933. A scrotal sac is not visible in this or any other of the photos or film taken, leading to the supposition that Benjamin was a female. However, photographic analysis in 2011 suggested that Benjamin was male.

The last captive thylacine, often referred to as Benjamin, lived as an endling; (the known last of its species), at Hobart Zoo until its death on the night of September 6, 1936.[114] Its source has long been disputed. Until recently, Elias Churchill was regularly quoted as being the captor, but there appears to be little evidence to support this claim. Two more recent candidates are far better placed evidentially as the probable source – the Kaine capture near Preolenna in 1931[115] and the Delphin capture near Waratah in 1930.[116] The thylacine died on the night of 6–7 September 1936. It is believed to have died as the result of neglect—locked out of its sheltered sleeping quarters, it was exposed to a rare occurrence of extreme Tasmanian weather: extreme heat during the day and freezing temperatures at night.[117] This thylacine features in the last known motion picture footage of a living specimen: 45 seconds of black-and-white footage showing the thylacine in its enclosure in a clip taken in 1933, by naturalist David Fleay.[118] In the film footage, the thylacine is seen seated, walking around the perimeter of its enclosure, yawning, sniffing the air, scratching itself (in the same manner as a dog), and lying down. Fleay was bitten on the buttock whilst shooting the film.[118] In 2021, a digitally colourised 80-second clip of Fleay's footage of Benjamin was released by the National Film and Sound Archive of Australia, to mark National Threatened Species Day. The digital colourisation process was completed by a Paris-based company, based on historic primary and secondary descriptions to ensure an as exact colour match as possible.[119][120]

Frank Darby, who claimed to have been a keeper at Hobart Zoo, suggested Benjamin as having been the animal's pet name in a newspaper article of May 1968. No documentation exists to suggest that it ever had a pet name, and Alison Reid (de facto curator at the zoo) and Michael Sharland (publicist for the zoo) denied that Frank Darby had ever worked at the zoo or that the name Benjamin was ever used for the animal. Darby also appears to be the source for the claim that the last thylacine was a male.[121] Robert Paddle was unable to uncover any records of any Frank Darby having been employed by Beaumaris/Hobart Zoo during the time that Reid or her father was in charge and noted several inconsistencies in the story Darby told during his interview in 1968.

The sex of the last captive thylacine has been a point of debate since its death at the Beaumaris Zoo in Hobart, Tasmania. In 2011, a detailed examination of a single frame from the motion film footage confirmed that the thylacine was male. When frame III is enlarged the scrotum can be seen; and by enhancing the frame, the outline of the individual testes is discernable.[122]

After the thylacine's death, the zoo expected that it would soon find a replacement,[109] and "Benjamin"'s death was not reported on in the media at the time.[123] Although there had been a conservation movement pressing for the thylacine's protection since 1901, driven in part by the increasing difficulty in obtaining specimens for overseas collections, political difficulties prevented any form of protection coming into force until 1936. Official protection of the species by the Tasmanian government was introduced on 10 July 1936, 59 days before the last known specimen died in captivity.[124]

A thylacine was reportedly shot and photographed at Mawbanna in 1938. A 1957 sighting from a helicopter could not be confirmed on the ground. An animal killed in Sandy Cape at night in 1961 was tentatively identified as a thylacine.[109] The results of subsequent searches indicated a strong possibility of the survival of the species in Tasmania into the 1960s. Searches by Dr. Eric Guiler and David Fleay in the northwest of Tasmania found footprints and scats that may have belonged to the animal, heard vocalisations matching the description of those of the thylacine, and collected anecdotal evidence from people reported having sighted the animal.

Despite the searches, no conclusive evidence was found to point to its continued existence in the wild.[37] Between 1967 and 1973, zoologist Jeremy Griffith and dairy farmer James Malley conducted what is regarded as the most intensive search ever carried out, including exhaustive surveys along Tasmania's west coast, installation of automatic camera stations, prompt investigations of claimed sightings, and in 1972 the creation of the Thylacine Expeditionary Research Team with Dr. Bob Brown, which concluded without finding any evidence of the thylacine's existence.[125]

The thylacine held the status of endangered species until the 1980s. International standards at the time stated that an animal could not be declared extinct until 50 years had passed without a confirmed record. Since no definitive proof of the thylacine's existence in the wild had been obtained for more than 50 years, it met that official criterion and was declared extinct by the International Union for Conservation of Nature in 1982[4] and by the Tasmanian government in 1986. The species was removed from Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) in 2013.[126]

Unconfirmed sightings
Map showing the location of reported sightings between 1936 and 1980 in Tasmania. Black = 1 reported sighting, red = 5 reported sightings.

The Department of Conservation and Land Management recorded 203 reports of sightings of the thylacine in Western Australia from 1936 to 1998.[67] On the mainland, sightings are most frequently reported in Southern Victoria.[127]

Map of reported sightings in the southwest of Western Australia

In 1982, a researcher with the Tasmania Parks and Wildlife Service, Hans Naarding, observed what he believed to be a thylacine for three minutes during the night at a site near Arthur River in northwestern Tasmania. The sighting led to an extensive year-long government-funded search.[128] In 1985, Aboriginal tracker Kevin Cameron produced five photographs which appear to show a digging thylacine, which he stated he took in Western Australia.[129]

In January 1995, a Parks and Wildlife officer reported observing a thylacine in the Pyengana region of northeastern Tasmania in the early hours of the morning. Later searches revealed no trace of the animal.[130] In 1997, it was reported that locals and missionaries near Mount Carstensz in Western New Guinea had sighted thylacines.[131][132] The locals had apparently known about them for many years but had not made an official report.[133] In February 2005 Klaus Emmerichs, a German tourist, claimed to have taken digital photographs of a thylacine he saw near the Lake St Clair National Park, but the authenticity of the photographs has not been established.[134] The photos were published in April 2006, fourteen months after the sighting. The photographs, which showed only the back of the animal, were said by those who studied them to be inconclusive as evidence of the thylacine's continued existence.[135]

In light of two detailed sightings around 1983 from the remote Cape York Peninsula of mainland Australia, scientists led by Bill Laurance announced plans in 2017 to survey the area for thylacines using camera traps.[136][137]

In 2017, 580 camera traps were deployed in North Queensland by James Cook University after two people – an experienced outdoorsman and a former Park Ranger – reported having seen a thylacine there in the 1980s but being too embarrassed to tell anyone at the time.[138][139]

According to the Department of Primary Industries, Parks, Water and Environment, there have been eight unconfirmed thylacine sighting reports between 2016 and 2019, with the latest unconfirmed visual sighting on 25 February 2018.[140]

Since the disappearance and effective extinction of the thylacine, speculation and searches for a living specimen have become a topic of interest to some members of the cryptozoology subculture.[141] The search for the animal has been the subject of books and articles, with many reported sightings that are largely regarded as dubious. According to writer Errol Fuller, the most likely record of the species persistence was proposed by Athol Douglas in the journal Cryptozoology, where Douglas challenges the carbon dating of the specimen found at Mundrabilla in South Australia as 4,500 years old; Douglas proposed instead that the well-preserved thylacine carcass was several months old when discovered. The dating of the specimen has not been reassessed.[142]

A preliminary 2021 study published by Brook et al.. compiles many of the alleged sightings of thylacines in Tasmania throughout the 20th century and claims that contrary to beliefs that thylacines went extinct in the 1930s, the Tasmanian thylacine may have actually lasted throughout the 20th century with a window of extinction between the 1980s and the present day, with the likely extinction date being between the late 1990s and early 2000s. The study claims that the reason behind the apparent lack of confirmed sightings during this wide and relatively recent time range is the lack of wide deployment of camera traps (which have been used to rediscover other elusive carnivores, such as the Zanzibar leopard) in Tasmania until the early 21st century, by which time the thylacine would be extinct or very nearly so. (This study has not yet been peer-reviewed.)[143][144]

Rewards

In 1983, the American media mogul Ted Turner offered a $100,000 reward for proof of the continued existence of the thylacine.[145] A letter sent in response to an inquiry by a thylacine-searcher, Murray McAllister in 2000, indicated that the offer had been withdrawn.[146] In March 2005, Australian news magazine The Bulletin, as part of its 125th anniversary celebrations, offered a $1.25 million reward for the safe capture of a live thylacine. When the offer closed at the end of June 2005, no one had produced any evidence of the animal's existence. An offer of $1.75 million has subsequently been offered by a Tasmanian tour operator, Stewart Malcolm.[135] Trapping is illegal under the terms of the thylacine's protection, so any reward made for its capture is invalid, since a trapping license would not be issued.[145]

Research
Specimen in the Oslo museum, showing colouration

The Australian Museum in Sydney began a cloning project in 1999.[147] The goal was to use genetic material from specimens taken and preserved in the early 20th century to clone new individuals and restore the species from extinction. Several molecular biologists have dismissed the project as a public relations stunt and its chief proponent, Mike Archer, received a 2002 nomination for the Australian Skeptics Bent Spoon Award for "the perpetrator of the most preposterous piece of paranormal or pseudo-scientific piffle".[148]

Skeleton in Cambridge University Museum of Zoology, England

In late 2002, the researchers had some success as they were able to extract replicable DNA from the specimens.[149] On 15 February 2005, the museum announced that it was stopping the project after tests showed the DNA retrieved from the specimens had been too badly degraded to be usable.[150][151] In May 2005, Archer, the University of New South Wales Dean of Science at the time, former director of the Australian Museum and evolutionary biologist, announced that the project was being restarted by a group of interested universities and a research institute.[135][152]

In 2008, researchers Andrew J. Pask and Marilyn B. Renfree from the University of Melbourne and Richard R. Behringer from the University of Texas at Austin reported that they managed to restore functionality of a gene Col2A1 enhancer obtained from 100-year-old ethanol-fixed thylacine tissues from museum collections. The genetic material was found working in transgenic mice. The research enhanced hopes of eventually restoring the population of thylacines.[153][154] That same year, another group of researchers successfully sequenced the complete thylacine mitochondrial genome from two museum specimens. Their success suggests that it may be feasible to sequence the complete thylacine nuclear genome from museum specimens. Their results were published in the journal Genome Research in 2009.[39]

Preserved thylacine pouch young specimens. Specimen f was found to belong to a different species of marsupial (quoll or Tasmanian devil).[72]

Mike Archer reported about the possibilities of resurrecting the thylacine and the gastric-brooding frog at TED2013.[155] Stewart Brand spoke at TED2013 about the ethics and possibilities of de-extinction, and made reference to thylacine in his talk.[156] A draft genome sequence of the thylacine was produced by Feigin et al. (2017) using the DNA extracted from an ethanol-preserved pouch young specimen provided by Museums Victoria. The neonatal development of the thylacine was also reconstructed from preserved pouch young specimens from several museum collections.[157] Researchers used the genome to study aspects of the thylacine's evolution and natural history, including the genetic basis of its convergence with canids, clarifying its evolutionary relationships with other marsupials and examining changes in its population size over time.[158] The genomic basis of the convergent evolution between the thylacine and grey wolf was further investigated in 2019,[159] with researchers identifying many non-coding genomic regions displaying accelerated rates of evolution, a test for genetic regions evolving under Positive Selection. In 2021,[160] researchers further identified a link between the convergent skull shapes of the thylacine and wolf,[158] and the previously identified genetic candidates.[159] It was reported that specific groups of skull bones, which develop from a common population of stem cells called Neural crest cells, showed strong similarity between the thylacine and wolf[160] and corresponded with the underlying convergent genetic candidates which influence these cells during development.[159]

Also in 2017, a reference library of 159 micrographic images of thylacine hair was jointly produced by CSIRO and Where Light Meets Dark, using scanning electron microscopy, metal-coated scanning electron microscopy, confocal laser scanning microscopy and optical light microscopy.[161] In 2018, Rehberg published a study into the appearance of thylacine stripes using infrared flash camera trap photography.[162]

Research into thylacines relies heavily on specimens held in museums and other institutions across the world. The number and distribution of these specimens has been recorded in the International Thylacine Specimen Database. As of 2022, 756 specimens are held in 115 museums and university collections in 23 countries.[163]

In August 2022, it was announced that the University of Melbourne will partner with Texas-based biotechnology company Colossal Biosciences to attempt to re-create the thylacine and return it to Tasmania.[164] The university had recently sequenced the genome of a juvenile thylacine specimen and is establishing a thylacine genetic restoration laboratory.[165][166][167][168][169]

Cultural significance

In Aboriginal tradition

Few Tasmanian Aboriginal beliefs about the thylacine have been recorded and preserved. One Nuenonne myth recorded by Jackson Cotton tells of a thylacine pup saving Palana, a spirit boy, from an attack by a giant kangaroo. Palana marked the pup's back with ochre as a mark of its bravery, giving thylacines their stripes.[170] A constellation, "Wurrawana Corinna" (identified as within or near Gemini), was also created as a commemoration of this mythic act of bravery.[171][172]

George Augustus Robinson recorded that, while some Aboriginal groups on Tasmania ate thylacines, others revered the animal and refused to do so.

The Kunwinjku on mainland Australia have preserved both a name for the thylacine (djankerrk) and an account of its behaviour. A Kunwinjku story tells of two ancestral thylacines hunting a kangaroo by biting at its tail, and the animals later falling from a cliff into a creek and transforming into fish. The thylacines transformed into archerfish, hence archerfish have stripes on their tails. Thylacines are also associated with waterfalls and the Rainbow Serpent.[173]

Akngwelye, an ancestral being from Arrernte Dreaming stories, is traditionally interpreted as a dingo, although there is some consensus that the animal was a thylacine in original tellings of the story.[174]

As a symbol
John Gould's lithographic plate from The Mammals of Australia
The Tasmanian coat of arms features thylacines as supporters.

The best known illustrations of Thylacinus cynocephalus were those in John Gould's The Mammals of Australia (1845–63), often copied since its publication and the most frequently reproduced,[175] and given further exposure by Cascade Brewery's appropriation for its label in 1987.[176] The government of Tasmania published a monochromatic reproduction of the same image in 1934,[177] the author Louisa Anne Meredith also copied it for Tasmanian Friends and Foes (1881).[175]

The thylacine has been used extensively as a symbol of Tasmania. The animal is featured on the official Tasmanian coat of arms.[178] It is used in the official logos for the Tasmanian government and the City of Launceston.[178] It is also used on the University of Tasmania's ceremonial mace and the badge of the submarine HMAS Dechaineux.[178] Since 1998, it has been prominently displayed on Tasmanian vehicle number plates.

Commemoration

Since 1996,[179] 7 September (the date in 1936 on which the last known thylacine died) has been commemorated in Australia as National Threatened Species Day.[180]

The plight of the thylacine was featured in a campaign for The Wilderness Society entitled We used to hunt thylacines.

Video games

In video games, boomerang-wielding Ty the Tasmanian Tiger is the star of his own trilogy during the 2000s.[181]

Tiny Tiger, a villain in the popular Crash Bandicoot video game series is a mutated thylacine.[182]

In Valorant, agent Skye has the ability to use a Tasmanian tiger to scout out enemies and clear bomb planting sites.[183]

Sports

The thylacine is the mascot for the Tasmanian cricket team.[184]

Movies and television

Characters in the early 1990s cartoon Taz-Mania included the neurotic Wendell T. Wolf, the last surviving Tasmanian wolf.[185]

The Hunter is a 2011 Australian drama film, based on the 1999 novel of the same name by Julia Leigh.[186] It stars Willem Dafoe, who plays a man hired to track down the Tasmanian tiger.

In the 2021 film, Extinct, a thylacine named Burnie, along with a group of other extinct animals, help the movie's main characters travel through time to rescue their species from extinction.[187]

Other

The thylacine has appeared in postage stamps from Australia, Equatorial Guinea, and Micronesia.[188]

See also
  • De-extinction / Breeding back
  • Fauna of Australia
  • List of extinct animals of Australia
  • Savage River National Park

References

Notes
  1. The photograph may even have involved photo manipulation.[112][113]

Citations
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Bibliography
  • Paddle, Robert (2000). The Last Tasmanian Tiger: the History and Extinction of the Thylacine. Cambridge University Press. ISBN 978-0-521-53154-2.

Further reading
  • Bailey, C. (2013) Shadow of the Thylacine. Five mile press. ISBN 978-1-74346-485-4
  • Freeman, Carol (2014). Paper Tiger: How Pictures Shaped the Thylacine (illustrated ed.). Hobart, Tasmania: Forty South Publishing. ISBN 978-0992279172.
  • Guiler, E. (1985) Thylacine: The Tragedy of the Tasmanian Tiger. Oxford University Press. ISBN 978-0-19-554603-3
  • Guiler, E. & Godard, P. (1998) Tasmanian Tiger: A Lesson to Be Learnt. Abrolhos Publishing. ISBN 978-0-9585791-0-0
  • Guiler, E. R. (1961a). "Breeding season of the thylacine". Journal of Mammalogy. 42 (3): 396–397. doi:10.2307/1377040. JSTOR 1377040.
  • Guiler, E. R. (1961b). "The former distribution and decline of the Thylacine". Australian Journal of Science. 23 (7): 207–210.
  • Heath, A. R. (2014) Thylacine: Confirming Tasmanian Tigers Still Live. Vivid Publishing. ISBN 9781925209402.
  • Leigh, J. (1999) The Hunter. Faber and Faber. ISBN 9780571200191.
  • Lord, C. (1927). "Existing Tasmanian marsupials". Papers and Proceedings of the Royal Society of Tasmania. Royal Society of Tasmania. 61: 17–24.
  • Lowry, D. C. (1967) "Discovery of a Thylacine (Tasmanian Tiger) Carcase in a Cave near Eucla, Western Australia". Helictite.
  • Owen, David (2003). Thylacine: the Tragic Tale of the Tasmanian Tiger. Allen & Unwin. ISBN 978-1-86508-758-0. Archived from the original on 1 April 2012. Retrieved 28 August 2010.
  • Pearce, R (1976). "Thylacines in Tasmania". Australian Mammal Society Bulletin. 3: 58.
  • Sleightholme, S. & Ayliffe, N. (2005). International Thylacine Specimen Database. CD-ROM. Master Copy: Zoological Society, London
  • Smith, S. J. (1980). "The Tasmanian Tiger – 1980. A report on an investigation of the current status of thylacine Thylacinus cynocephalus, funded by the World Wildlife Fund Australia". Hobart: National Parks and Wildlife Service, Tasmania.

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