Tasmanian devil

Karyotype of male Tasmanian devil.

The Tasmanian devil's genome was sequenced in 2010 by the Wellcome Trust Sanger Institute.[21] Like all dasyurids, the devil has 14 chromosomes.[22] Devils have a low genetic diversity compared to other Australian marsupials and placental carnivores; this is consistent with a founder effect as allelic size ranges were low and nearly continuous throughout all subpopulations measured. Allelic diversity was measured at 2.7–3.3 in the subpopulations sampled, and heterozygosity was in the range 0.386–0.467.[23] According to a study by Menna Jones, "gene flow appears extensive up to 50 km (31 mi)", meaning a high assignment rate to source or close neighbour populations "in agreement with movement data. At larger scales (150–250 km or 90–200 mi), gene flow is reduced but there is no evidence for isolation by distance".[23] Island effects may also have contributed to their low genetic diversity. Periods of low population density may also have created moderate population bottlenecks, reducing genetic diversity.[23] Low genetic diversity is thought to have been a feature in the Tasmanian devil population since the mid-Holocene.[24] Outbreaks of devil facial tumour disease (DFTD) cause an increase in inbreeding.[25] A sub-population of devils in the north-west of the state is genetically distinct from other devils,[26] but there is some exchange between the two groups.[27]

One strand conformation polymorphism analysis (OSCP) on the major histocompatibility complex (MHC) class I domain taken from various locations across Tasmania showed 25 different types, and showed a different pattern of MHC types in north-western Tasmania to eastern Tasmania. Those devils in the east of the state have less MHC diversity; 30% are of the same type as the tumour (type 1), and 24% are of type A.[28] Seven of every ten devils in the east are of type A, D, G or 1, which are linked to DFTD; whereas only 55% of the western devils fall into these MHC categories. Of the 25 MHC types, 40% are exclusive to the western devils. Although the north-west population is less genetically diverse overall, it has higher MHC gene diversity, which allows them to mount an immune response to DFTD. According to this research, mixing the devils may increase the chance of disease.[28] Of the fifteen different regions in Tasmania surveyed in this research, six were in the eastern half of the island. In the eastern half, Epping Forest had only two different types, 75% being type O. In the Buckland-Nugent area, only three types were present, and there were an average of 5.33 different types per location. In contrast, in the west, Cape Sorell yielded three types, and Togari North-Christmas Hills yielded six, but the other seven sites all had at least eight MHC types, and West Pencil Pine had 15 types. There was an average of 10.11 MHC types per site in the west.[28] Recent research has suggested that the wild population of devils are rapidly evolving a resistance to DFTD.[29]

A devil eating roadkill

Tasmanian devils can take prey up to the size of a small kangaroo, but in practice they are opportunistic and eat carrion more often than they hunt live prey. Although the devil favours wombats because of the ease of predation and high fat content, it will eat all small native mammals such as wallabies,[78] bettong and potoroos, domestic mammals (including sheep and rabbits),[78] birds (including penguins),[79] fish, fruit, vegetable matter, insects, tadpoles, frogs and reptiles. Their diet is widely varied and depends on the food available.[37][80][81][82] Before the extinction of the thylacine, the Tasmanian devil ate thylacine joeys left alone in dens when their parents were away. This may have helped to hasten the extinction of the thylacine, which also ate devils.[47] They are known to hunt water rats by the sea and forage on dead fish that have been washed ashore. Near human habitation, they can also steal shoes and chew on them,[80] and eat the legs of otherwise robust sheep that have slipped in wooden shearing sheds, leaving their legs dangling below.[62] Other unusual matter observed in devil scats includes collars and tags of devoured animals, intact echidna spines, pencil, plastic and jeans.[59] Devils can bite through metal traps, and tend to reserve their strong jaws for escaping captivity rather than breaking into food storage.[59] Due to their relative lack of speed, they can not run down a wallaby or a rabbit, but they can attack animals that have become slow due to illness.[80] They survey flocks of sheep by sniffing them from 10–15 m (33–49 ft) away and attack if the prey is ill. The sheep stamp their feet in a show of strength.[59]

Despite their lack of extreme speed, there have been reports that devils can run at 25 km/h (16 mph) for 1.5 km (0.93 mi), and it has been conjectured that, before European immigration and the introduction of livestock, vehicles and roadkill, they would have had to chase other native animals at a reasonable pace to find food.[62] Pemberton has reported that they can average 10 km/h (6.2 mph) for "extended periods" on several nights per week, and that they run for long distances before sitting still for up to half an hour, something that has been interpreted as evidence of ambush predation.[62]

Devils can dig to forage corpses, in one case digging down to eat the corpse of a buried horse that had died due to illness. They are known to eat animal cadavers by first ripping out the digestive system, which is the softest part of the anatomy, and they often reside in the resulting cavity while they are eating.[80]

On average, devils eat about 15% of their body weight each day, although they can eat up to 40% of their body weight in 30 minutes if the opportunity arises.[39] This means they can become very heavy and lethargic after a large meal; in this state they tend to waddle away slowly and lie down, becoming easy to approach. This has led to a belief that such eating habits became possible due to the lack of a predator to attack such bloated individuals.[81]

Tasmanian devils can eliminate all traces of a carcass of a smaller animal, devouring the bones and fur if desired.[83] In this respect, devils have earned the gratitude of Tasmanian farmers, as the speed at which they clean a carcass helps prevent the spread of insects that might otherwise harm livestock.[84] Some of these dead animals are disposed of when the devils haul off the excess feed back to their residence to continue eating at a later time.[80]

The diet of a devil can vary substantially for males and females, and seasonally, according to studies at Cradle Mountain. In winter, males prefer medium mammals over larger ones, with a ratio of 4:5, but in summer, they prefer larger prey in a 7:2 ratio. These two categories accounted for more than 95% of the diet. Females are less inclined to target large prey, but have the same seasonal bias. In winter, large and medium mammals account for 25% and 58% each, with 7% small mammals and 10% birds. In summer, the first two categories account for 61% and 37% respectively.[61]

Juvenile devils are sometimes known to climb trees;[85] in addition to small vertebrates and invertebrates, juveniles climb trees to eat grubs and birds' eggs.[60] Juveniles have also been observed climbing into nests and capturing birds.[64] Throughout the year, adult devils derive 16.2% of their biomass intake from arboreal species, almost all of which is possum meat, just 1.0% being large birds. From February to July, subadult devils derive 35.8% of their biomass intake from arboreal life, 12.2% being small birds and 23.2% being possums. Female devils in winter source 40.0% of their intake from arboreal species, including 26.7% from possums and 8.9% from various birds.[64] Not all of these animals were caught while they were in trees, but this high figure for females, which is higher than for male spotted-tailed quolls during the same season, is unusual, as the devil has inferior tree climbing skills.[64]

Three Tasmanian devils feeding. Eating is a social event for the Tasmanian devil, and groups of 2 to 5 are common.

Although they hunt alone,[37] there have been unsubstantiated claims of communal hunting, where one devil drives prey out of its habitat and an accomplice attacks.[80] Eating is a social event for the Tasmanian devil. This combination of a solitary animal that eats communally makes the devil unique among carnivores.[60] Much of the noise attributed to the animal is a result of raucous communal eating, at which up to 12 individuals can gather,[39] although groups of two to five are common;[86] it can often be heard several kilometres away. This has been interpreted as notifications to colleagues to share in the meal, so that food is not wasted by rot and energy is saved.[80] The amount of noise is correlated to the size of the carcass.[80] The devils eat in accordance with a system. Juveniles are active at dusk, so they tend to reach the source before the adults.[81] Typically, the dominant animal eats until it is satiated and leaves, fighting off any challengers in the meantime. Defeated animals run into the bush with their hair and tail erect, their conqueror in pursuit and biting their victim's rear where possible. Disputes are less common as the food source increases as the motive appears to be getting sufficient food rather than oppressing other devils.[81] When quolls are eating a carcass, devils will tend to chase them away.[64] This is a substantial problem for spotted-tailed quolls, as they kill relatively large possums and cannot finish their meal before devils arrive. In contrast, the smaller eastern quolls prey on much smaller victims, and can complete feeding before devils turn up.[64] This is seen as a possible reason for the relatively small population of spotted-tailed quolls.[64]

A study of feeding devils identified twenty physical postures, including their characteristic vicious yawn, and eleven different vocal sounds that devils use to communicate as they feed.[39] They usually establish dominance by sound and physical posturing,[87] although fighting does occur.[39] The white patches on the devil are visible to the night-vision of its colleagues.[81] Chemical gestures are also used.[81] Adult males are the most aggressive,[88] and scarring is common.[89] They can also stand on their hind legs and push each other's shoulders with their front legs and heads, similar to sumo wrestling.[81] Torn flesh around the mouth and teeth, as well as punctures in the rump, can sometimes be observed, although these can also be inflicted during breeding fights.[81]

Digestion is very fast in dasyurids and, for the Tasmanian devil, the few hours taken for food to pass through the small gut is a long period in comparison to some other dasyuridae.[90] Devils are known to return to the same places to defecate, and to do so at a communal location, called a devil latrine.[91] It is believed that the communal defecation may be a means of communication that is not well understood.[91] Devil scats are very large compared to body size; they are on average 15 centimetres (5.9 in) long, but there have been samples that are 25 centimetres (9.8 in) in length.[91] They are characteristically grey in colour due to digested bones, or have bone fragments included.[26]

Owen and Pemberton believe that the relationship between Tasmanian devils and thylacines was "close and complex", as they competed directly for prey and probably also for shelter. The thylacines preyed on the devils, the devils scavenged from the thylacine's kills, and the devils ate thylacine young. Menna Jones hypothesises that the two species shared the role of apex predator in Tasmania.[92] Wedge-tailed eagles have a similar carrion-based diet to the devils and are regarded as competitors.[93] Quolls and devils are also seen as being in direct competition in Tasmania. Jones believed that the quoll has evolved into its current state in just 100–200 generations of around two years as determined by the equal spacing effect on the devil, the largest species, the spotted-tail quoll, and the smallest species, the eastern quoll.[94] Both the Tasmanian devil and the quolls appears to have evolved up to 50 times faster than the average evolutionary rate amongst mammals.[95]

Developmental steps in the maturation of Tasmanian devil young. The diagonal lines indicate the amount of time the changes take; for example, it takes about 90 days for a devil to develop fur over all its body.

Females start to breed when they reach sexual maturity, typically in their second year. At this point, they become fertile once a year, producing multiple ova while in heat.[96] As prey is most abundant in spring and early summer, the devil's reproductive cycle starts in March or April so that the end of the weaning period coincides with the maximisation of food supplies in the wild for the newly roaming young devils.[97]

Occurring in March, mating takes places in sheltered locations during both day and night. Males fight over females in the breeding season, and female devils will mate with the dominant male.[37][98] Females can ovulate up to three times in a 21-day period, and copulation can take five days; one instance of a couple being in the mating den for eight days has been recorded.[98] Devils are not monogamous, and females will mate with several males if not guarded after mating; males also reproduce with several females during a season.[37][98] Females have been shown to be selective in an attempt to ensure the best genetic offspring,[98] for example, fighting off the advances of smaller males.[27] Males often keep their mates in custody in the den, or take them along if they need to drink, lest they engage in infidelity.[98]

Males can produce up to 16 offspring over their lifetime, while females average four mating seasons and 12 offspring.[98] Theoretically this means that a devil population can double on an annual basis and make the species insulated against high mortality.[99] The pregnancy rate is high; 80% of two-year-old females were observed with newborns in their pouches during the mating season.[98] More recent studies of breeding place the mating season between February and June, as opposed to between February and March.[26]

Gestation lasts 21 days, and devils give birth to 20–30 young standing up,[37][98] each weighing approximately 0.18–0.24 grams (0.0063–0.0085 oz).[55] Embryonic diapause does not occur.[96] At birth, the front limb has well-developed digits with claws; unlike many marsupials, the claws of baby devils are not deciduous. As with most other marsupials, the forelimb is longer (0.26–0.43 cm or 0.10–0.17 in) than the rear limb (0.20–0.28 cm or 0.079–0.110 in), the eyes are spots, and the body is pink. There are no external ears or openings. Unusually, the sex can be determined at birth, with an external scrotum present.[96]

Tasmanian devil young are variously called "pups",[37] "joeys",[100] or "imps".[101] When the young are born, competition is fierce as they move from the vagina in a sticky flow of mucus to the pouch. Once inside the pouch, they each remain attached to a nipple for the next 100 days. The female Tasmanian devil's pouch, like that of the wombat, opens to the rear, so it is physically difficult for the female to interact with young inside the pouch. Despite the large litter at birth, the female has only four nipples, so there are never more than four babies nursing in the pouch, and the older a female devil gets, the smaller her litters will become. Once the young have made contact with the nipple, it expands, resulting in the oversized nipple being firmly clamped inside the newborn and ensuring that the newborn does not fall out of the pouch.[37][98] On average, more females survive than males,[96] and up to 60% of young do not survive to maturity.[60] Milk replacements are often used for devils that have been bred in captivity, for orphaned devils or young who are born to diseased mothers. Little is known about the composition of the devil's milk compared to other marsupials.[102]

Inside the pouch, the nourished young develop quickly. In the second week, the rhinarium becomes distinctive and heavily pigmented.[96] At 15 days, the external parts of the ear are visible, although these are attached to the head and do not open out until the devil is around 10 weeks old. The ear begins blackening after around 40 days, when it is less than 1 cm (0.39 in) long, and by the time the ear becomes erect, it is between 1.2 and 1.6 cm (0.47 and 0.63 in). Eyelids are apparent at 16 days, whiskers at 17 days, and the lips at 20 days.[96] The devils can make squeaking noises after eight weeks, and after around 10–11 weeks, the lips can open.[96] Despite the formation of eyelids, they do not open for three months, although eyelashes form at around 50 days.[96] The young—up to this point they are pink—start to grow fur at 49 days and have a full coat by 90 days. The fur growing process starts at the snout and proceeds back through the body, although the tail attains fur before the rump, which is the last part of the body to become covered. Just before the start of the furring process, the colour of the bare devil's skin will darken and become black or dark grey in the tail.[96]

Three young devils sunbathing

The devils have a complete set of facial vibrissae and ulnar carpels, although it is devoid of anconeal vibrissae. During the third week, the mystacials and ulnarcarpals are the first to form. Subsequently, the infraorbital, interramal, supraorbital and submental vibrissae form. The last four typically occur between the 26th and 39th day.[96] Their eyes open shortly after their fur coat develops—between 87 and 93 days—and their mouths can relax their hold of the nipple at 100 days.[96] They leave the pouch 105 days after birth, appearing as small copies of the parent and weighing around 200 grams (7.1 oz).[96] Zoologist Eric Guiler recorded its size at this time as follows: a crown-snout length of 5.87 cm (2.31 in), tail length of 5.78 cm (2.28 in), pes length 2.94 cm (1.16 in), manus 2.30 cm (0.91 in), shank 4.16 cm (1.64 in), forearm 4.34 cm (1.71 in) and crown-rump length is 11.9 cm (4.7 in).[96] During this period, the devils lengthen at a roughly linear rate.[96]

After being ejected, the devils stay outside the pouch, but they remain in the den for around another three months, first venturing outside the den between October and December before becoming independent in January. During this transitional phase out of the pouch, the young devils are relatively safe from predation as they are generally accompanied. When the mother is hunting they can stay inside a shelter or come along, often riding on their mother's back. During this time they continue to drink their mother's milk. Female devils are occupied with raising their young for all but approximately six weeks of the year.[96][103] The milk contains a higher amount of iron than the milk of placental mammals.[27] In Guiler's 1970 study, no females died while rearing their offspring in the pouch. After leaving the pouch, the devils grow by around 0.5 kg (1.1 lb) a month until they are six months old.[96] While most pups will survive to be weaned,[26] Guiler reported that up to three fifths of devils do not reach maturity.[60] As juveniles are more crepuscular than adults, their appearance in the open during summer gives the impression to humans of a population boom.[60] A study into the success of translocated devils that were orphaned and raised in captivity found that young devils who had consistently engaged with new experiences while they were in captivity survived better than young who had not.[104]

At least two major population declines, possibly due to disease epidemics, have occurred in recorded history: in 1909 and 1950.[30] The devil was also reported as scarce in the 1850s.[115] It is difficult to estimate the size of the devil population.[116] In the mid-1990s, the population was estimated at 130,000–150,000 animals,[26] but this is likely to have been an overestimate.[116] The Tasmanian devil's population has been calculated in 2008 by Tasmania's Department of Primary Industries and Water as being in the range of 10,000 to 100,000 individuals, with 20,000 to 50,000 mature individuals being likely.[37] Experts estimate that the devil has suffered a more than 80% decline in its population since the mid-1990s and that only around 10,000–15,000 remain in the wild as of 2008.[117]

The species was listed as vulnerable under the Tasmanian Threatened Species Protection Act 1995 in 2005[118] and the Australian Environment Protection and Biodiversity Conservation Act 1999[26] in 2006, which means that it is at risk of extinction in the "medium term".[50] The IUCN classified the Tasmanian devil in the lower risk/least concern category in 1996, but in 2009 they reclassified it as endangered.[1] Appropriate wildlife refuges such as Savage River National Park in North West Tasmania provide hope for their survival.

The first European Tasmanian settlers ate Tasmanian devil, which they described as tasting like veal.[119] As it was believed devils would hunt and kill livestock, possibly due to strong imagery of packs of devils eating weak sheep, a bounty scheme to remove the devil from rural properties was introduced as early as 1830.[120] However, Guiler's research contended that the real cause of livestock losses was poor land management policies and feral dogs.[120] In areas where the devil is now absent, poultry has continued to be killed by quolls. In earlier times, hunting possums and wallabies for fur was a big business—more than 900,000 animals were hunted in 1923—and this resulted in a continuation of bounty hunting of devils as they were thought to be a major threat to the fur industry, even though quolls were more adept at hunting the animals in question.[121] Over the next 100 years, trapping and poisoning[122] brought them to the brink of extinction.[111]

After the death of the last thylacine in 1936,[123] the Tasmanian devil was protected by law in June 1941 and the population slowly recovered.[111] In the 1950s, with reports of increasing numbers, some permits to capture devils were granted after complaints of livestock damage. In 1966, poisoning permits were issued although attempts to have the animal unprotected failed.[124] During this time environmentalists also became more outspoken, particularly as scientific studies provided new data suggesting the threat of devils to livestock had been vastly exaggerated.[125] Numbers may have peaked in the early 1970s after a population boom; in 1975 they were reported to be lower, possibly due to overpopulation and consequent lack of food.[126] Another report of overpopulation and livestock damage was reported in 1987.[127] The following year, Trichinella spiralis, a parasite which kills animals and can infect humans, was found in devils and minor panic broke out before scientists assured the public that 30% of devils had it but that they could not transmit it to other species.[128] Control permits were ended in the 1990s, but illegal killing continues to a limited extent, albeit "locally intense". This is not considered a substantial problem for the survival of the devil.[50] Approximately 10,000 devils were killed per year in the mid-1990s.[27] A selective culling program has taken place to remove individuals affected with DFTD, and has been shown to not slow the rate of disease progression or reduced the number of animals dying.[129] A model has been tested to find out whether culling devils infected with DFTD would assist in the survival of the species, and it has found that culling would not be a suitable strategy to employ.[130]

A road sign telling drivers that there may be devils nearby

Motor vehicles are a threat to localised populations of non-abundant Tasmanian mammals,[131][132] and a 2010 study showed that devils were particularly vulnerable. A study of nine species, mostly marsupials of a similar size, showed that devils were more difficult for drivers to detect and avoid. At high beam, devils had the lowest detection distance, 40% closer than the median. This requires a 20% reduction in speed for a motorist to avoid the devil. For low beam, the devils had the second shortest detection distance, 16% below the median. For avoidance of roadkill to be feasible, motorists would have to drive at around half the current speed limit in rural areas.[131] A study in the 1990s on a localised population of devils in a national park in Tasmania recorded a halving of the population after a hitherto gravel access road was upgraded, surfaced with bitumen and widened. At the same time, there was a large increase in deaths caused by vehicles along the new road; there had been none in the preceding six months.[132]

The vast majority of deaths occurred in the sealed portion of the road, believed to be due to an increase in speeds.[132] It was also conjectured that the animals were harder to see against the dark bitumen instead of the light gravel. The devil and quoll are especially vulnerable as they often try to retrieve roadkill for food and travel along the road. To alleviate the problem, traffic slowing measures, man-made pathways that offer alternative routes for devils, education campaigns, and the installation of light reflectors to indicate oncoming vehicles have been implemented. They are credited with decreases in roadkill.[132] Devils have often been victims of roadkill when they are retrieving other roadkill. Work by scientist Menna Jones and a group of conservation volunteers to remove dead animals from the road resulted in a significant reduction in devil traffic deaths.[84] It was estimated that 3,392 devils, or between 3.8 and 5.7% of the population, were being killed annually by vehicles in 2001–04.[50] In 2009, the Save the Tasmanian Devil group launched the "Roadkill Project", which allowed members of the public to report sightings of devils which had been killed on the road.[133] On 25 September 2015, 20 immunised devils were microchipped and released in Narawntapu National Park. By 5 October 4 had been hit by cars, prompting Samantha Fox, leader of Save the Tasmanian Devil, to describe roadkill as being the biggest threat to the Tasmanian devil after DFTD.[134] A series of solar-powered alarms have been trialled that make noises and flash lights when cars are approaching, warning the animals. The trial ran for 18 months and the trial area had two-thirds less deaths than the control.[135][136]

Devil facial tumour disease causes tumours to form in and around the mouth, interfering with feeding and eventually leading to death by starvation

First seen in 1996 in Mount William in northeastern Tasmania, devil facial tumour disease (DFTD) has ravaged Tasmania's wild devils, and estimates of the impact range from 20% to as much as an 80% decline in the devil population, with over 65% of the state affected. The state's west coast area and far north-west are the only places where devils are tumour free.[137][138][139] Individual devils die within months of infection.[140] The disease is an example of transmissible cancer, which means that it is contagious and passed from one animal to another.[141] This tumour is able to pass between hosts without inducing a response from the host's immune system.[142] Dominant devils who engage in more biting behaviour are more exposed to the disease.[143]

Wild Tasmanian devil populations are being monitored to track the spread of the disease and to identify changes in disease prevalence. Field monitoring involves trapping devils within a defined area to check for the presence of the disease and determine the number of affected animals. The same area is visited repeatedly to characterise the spread of the disease over time. So far, it has been established that the short-term effects of the disease in an area can be severe. Long-term monitoring at replicated sites will be essential to assess whether these effects remain, or whether populations can recover.[139] Field workers are also testing the effectiveness of disease suppression by trapping and removing diseased devils. It is hoped that the removal of diseased devils from wild populations should decrease disease prevalence and allow more devils to survive beyond their juvenile years and breed.[139] In March 2017, scientists at the University of Tasmania presented an apparent first report of having successfully treated Tasmanian devils with the disease, by injecting live cancer cells into the infected devils to stimulate their immune system to recognise and fight the disease.[144]

At Healesville Sanctuary, Victoria

Early attempts to breed Tasmanian devils in captivity had limited success. Mary Roberts bred a pair at Beaumaris Zoo (which she named Billy and Truganini) in 1913. However, although advised to remove Billy, Roberts found Truganini too distressed by his absence, and returned him. The first litter was presumed eaten by Billy, but a second litter in 1914 survived, after Billy was removed. Roberts wrote an article on keeping and breeding the devils for the London Zoological Society.[154] Even by 1934, successful breeding of the devil was rare.[157] In a study on the growth of young devils in captivity, some developmental stages were very different from those reported by Guiler. The pinnae were free on day 36, and eyes opened later, on days 115–121.[158] In general, females tend to retain more stress after being taken into captivity than males.[159]

A plan to create "insurance populations" of disease-free devils has been ongoing since 2005. As of June 2012, the insurance population has reached a combined total of 500 animals and representing over 98% of the genetic diversity of this species.[160] Most of these devils are living in Australian zoos and wildlife reserves. Beginning in November 2012 however, in an effort to create a population that is both wild and disease-free, Tasmanian devils have been relocated to Maria Island, a mountainous island off the east coast of Tasmania.[160] The Maria Island population has grown from a starting population of twenty-eight to 90, and experts will soon begin transferring healthy devils back to the mainland population.[161] A study on the survival rates of the Maria Island population found that in contrast to other carnivores raised in captivity, the Tasmanian devils were not adversely affected by being born in captivity when released on Maria Island.[162]

Tasmanian devils were displayed in various zoos around the world from the 1850s onwards.[163] In the 1950s several animals were given to European zoos.[164] In October 2005 the Tasmanian government sent four devils, two male and two female, to the Copenhagen Zoo, following the birth of the first son of Frederik, Crown Prince of Denmark and his Tasmanian-born wife Mary.[165] Due to restrictions on their export by the Australian government, at the time these were the only devils known to be living outside Australia.[26] In June 2013, due to the successes of the insurance population program, it was planned to send devils to other zoos around the world in a pilot program.[166] San Diego Zoo Wildlife Alliance and Albuquerque Biopark were selected to participate in the program,[167] and Wellington Zoo and Auckland Zoo soon followed.[168] In the United States, four additional zoos have since been selected as part of the Australian government's Save the Tasmanian Devil program, the zoos selected were: the Fort Wayne Children's Zoo,[169] the Los Angeles Zoo,[170] the Saint Louis Zoo,[171] and the Toledo Zoo.[172] Captive devils are usually forced to stay awake during the day to cater to visitors, rather than following their natural nocturnal style.[173]

There have been reports and suspicions of illegal trading in the past. In 1997, a devil turned up in Western Australia; it had not escaped from any licensed keeper. During the 1990s there were internet sites in the US that were advertising devil sales, and rumours that some US Navy personnel had tried to buy them illegally during a visit to Tasmania.[174]

Warner Bros' Tasmanian Devil "Taz" at a parade in California

The devil is an iconic animal within Australia, and particularly associated with Tasmania. The animal is used as the emblem of the Tasmanian National Parks and Wildlife Service,[37] and the former Tasmanian Australian rules football team which played in the Victorian Football League was known as the Devils.[175] The Hobart Devils were once part of the National Basketball League.[176] The devil has appeared on several commemorative coins in Australia over the years.[177][178] Cascade Brewery in Tasmania sells a ginger beer with a Tasmanian devil on the label.[179] In 2015, the Tasmanian devil was chosen as Tasmania's state emblem.[180]

Tasmanian devils are popular with tourists, and the director of the Tasmanian Devil Conservation Park has described their possible extinction as "a really significant blow for Australian and Tasmanian tourism".[181] There has also been a multimillion-dollar proposal to build a giant 19 m-high, 35 m-long devil in Launceston in northern Tasmania as a tourist attraction.[182] Devils began to be used as ecotourism in the 1970s, when studies showed that the animals were often the only things known about Tasmania overseas, and suggested that they should therefore be the centrepiece of marketing efforts, resulting in some devils being taken on promotional tours.[183]

The Tasmanian devil is probably best known internationally as the inspiration for the Looney Tunes cartoon character the Tasmanian Devil, or "Taz" in 1954. Little known at the time, the loud hyperactive cartoon character has little in common with the real life animal.[184] After a few shorts between 1957 and 1964, the character was retired until the 1990s, when he gained his own show, Taz-Mania, and again became popular.[185] In 1997, a newspaper report noted that Warner Bros. had "trademarked the character and registered the name Tasmanian Devil", and that this trademark "was policed", including an eight-year legal case to allow a Tasmanian company to call a fishing lure "Tasmanian Devil". Debate followed, and a delegation from the Tasmanian government met with Warner Bros.[186] Ray Groom, the Tourism Minister, later announced that a "verbal agreement" had been reached. An annual fee would be paid to Warner Bros. in return for the Government of Tasmania being able to use the image of Taz for "marketing purposes". This agreement later disappeared.[187] In 2006, Warner Bros. permitted the Government of Tasmania to sell stuffed toys of Taz with profits funnelled into research on DFTD.[188]