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{{Short description|Australian carnivorous marsupial}} {{other uses|Tasmanian Devil (disambiguation)}} {{Featured article}} {{Pp-vandalism|small=yes}} {{Use Australian English|date=July 2011}} {{Use dmy dates|date=April 2020}} {{Speciesbox | name = Tasmanian devil | fossil_range = [[Holocene]] | image = Sarcophilus harrisii taranna.jpg | image_caption = | image_alt = A Tasmanian devil | status = EN | status_system = IUCN3.1 | status_ref = <ref name="iucn">{{cite iucn |author=Hawkins, C.E. |author2=McCallum, H. |author3=Mooney, N. |author4=Jones, M. |author5=Holdsworth, M. |date=2008 |title=''Sarcophilus harrisii'' |volume=2008 |page=e.T40540A10331066 |doi=10.2305/IUCN.UK.2008.RLTS.T40540A10331066.en |access-date=19 November 2021}}</ref> | genus = Sarcophilus | species = harrisii | authority = ([[Pierre Boitard|Boitard]], 1841)<ref>{{cite book|last1=Boitard|first1=[Pierre]|title=Le Jardin des plantes: Description et mœurs des mammifères de la Ménagerie et du Muséum d'histoire naturelle|date=n.d.|publisher=Gustave Barba|location=Paris|page=204|chapter-url=https://www.biodiversitylibrary.org/page/9815348|chapter=L'Ursin de Harris}}</ref> | range_map = Sarcophilus harrisii extent.png | range_map_caption = Distribution of the Tasmanian devil on Tasmania in grey (note: reintroduced New South Wales distribution not mapped) | range_map_alt = A map showing one large island (Tasmania) and two small islands north of it. The whole of Tasmania is coloured in and the waters and small islands are not, as the devil is not extant there. }} The '''Tasmanian devil''' ('''''Sarcophilus harrisii'''''; [[palawa kani]]: ''purinina'')<ref>{{Cite journal |date=2023-12-01 |title=Takayna/Tarkine and the EPBC Act : From Heritage Frameworks to Habitat Thinking |url=https://www.taylorfrancis.com/chapters/edit/10.4324/9781003389569-3/takayna-tarkine-epbc-act-brad-jessup-christine-parker |journal=Taylor & Francis |language=en |doi=10.4324/9781003389569-3 |archive-url=http://web.archive.org/web/20240423073008/https://www.taylorfrancis.com/chapters/edit/10.4324/9781003389569-3/takayna-tarkine-epbc-act-brad-jessup-christine-parker |archive-date=2024-04-23}}</ref> is a carnivorous [[marsupial]] of the family [[Dasyuridae]]. It was formerly present across mainland Australia, but became extinct there around 3,500 years ago; it is now confined to the island of [[Tasmania]]. The size of a small dog, the Tasmanian devil became the [[Largest mammals#Marsupials (Marsupialia)|largest carnivorous marsupial]] in the world following the [[extinction]] of the [[thylacine]] in 1936. It is related to [[quolls]], and distantly related to the thylacine. It is characterised by its stocky and muscular build, black fur, pungent odour, extremely loud and disturbing screech, keen sense of smell, and ferocity when feeding. The Tasmanian devil's large head and neck allow it to generate among the strongest bites per unit body mass of any extant predatory land mammal. It hunts prey and scavenges on [[carrion]]. Although devils are usually solitary, they sometimes eat and defecate together in a communal location. Unlike most other [[Dasyuridae|dasyurids]], the devil [[thermoregulation|thermoregulates]] effectively, and is active during the middle of the day without overheating. Despite its rotund appearance, it is capable of surprising speed and endurance, and can climb trees and swim across rivers. Devils are not [[Monogamy in animals|monogamous]]. Males fight one another for females, and guard their partners to prevent female infidelity. Females can ovulate three times in as many weeks during the [[seasonal breeder|mating season]], and 80% of two-year-old females are seen to be pregnant during the annual mating season. Females average four breeding seasons in their life, and give birth to 20 to 30 live young after three weeks' gestation. The newborn are pink, lack fur, have indistinct facial features, and weigh around {{cvt|0.20|g}} at birth. As there are only four nipples in the pouch, competition is fierce, and few newborns survive. The young grow rapidly, and are ejected from the pouch after around 100 days, weighing roughly {{cvt|200|g}}. The young become independent after around nine months. In 1941, devils became officially protected. Since the late 1990s, the [[devil facial tumour disease]] (DFTD) has drastically reduced the population and now threatens the survival of the species, which in 2008 was declared to be [[endangered species|endangered]]. Starting in 2013, Tasmanian devils are again being sent to zoos around the world as part of the [[Australian government]]'s Save the Tasmanian Devil Program. The devil is an iconic symbol of Tasmania and many organisations, groups and products associated with the state use the animal in their logos. It is seen as an important attractor of tourists to Tasmania and has come to worldwide attention through the ''[[Looney Tunes]]'' character [[Tasmanian Devil (Looney Tunes)|of the same name]]. ==Taxonomy== Believing it to be a type of [[Ameridelphia|opossum]], naturalist [[George Prideaux Robert Harris|George Harris]] wrote the first published description of the Tasmanian devil in 1807, naming it ''Didelphis ursina'',<ref name=Harris1807/> due to its bearlike characteristics such as the round ear.<ref>Owen and Pemberton, p. 79.</ref> He had earlier made a presentation on the topic at the [[Zoological Society of London]].<ref name="Owen and Pemberton-8"/> However, that particular binomial name had been given to the [[common wombat]] (later reclassified as ''Vombatus ursinus'') by [[George Shaw (biologist)|George Shaw]] in 1800, and was hence unavailable.<ref name=MSW3>{{MSW3 Dasyuromorphia|id=10800047|page=28}}</ref> In 1838, a specimen was named ''Dasyurus laniarius'' by [[Richard Owen]],<ref name="Mitchell-1839">{{cite book |last=Mitchell |first=Thomas |author-link=Thomas Mitchell (explorer) |title=Three Expeditions into the Interior, Volume II |publisher=T. & W. Boone |year=1839}} [http://gutenberg.net.au/ebooks/e00036.html Online] at Project Gutenberg Australia.</ref> but by 1877 he had relegated it to ''Sarcophilus''. The modern Tasmanian devil was named ''Sarcophilus harrisii'' ("Harris's flesh-lover") by French naturalist [[Pierre Boitard]] in 1841.<ref>{{cite journal|last1=Stephenson|first1=N. G.|title=Growth gradients among fossil monotremes and marsupials {{!}} The Palaeontological Association|journal=Palaeontology|date=1963|volume=6|issue=4|pages=615–624|url=http://www.palass.org/publications/palaeontology-journal/archive/6/4/article_pp615-624|language=en}}</ref> A later revision of the devil's taxonomy, published in 1987, attempted to change the species name to ''[[Sarcophilus laniarius]]'' based on mainland fossil records of only a few animals.<ref name=Werdelin1987/> However, this was not accepted by the taxonomic community at large; the name ''S. harrisii'' has been retained and ''S. laniarius'' relegated to a fossil species.<ref name=MSW3/> "[[Beelzebub]]'s pup" was an early vernacular name given to it by the explorers of Tasmania, in reference to a religious figure who is a prince of hell and an assistant of [[Satan]];<ref name="Owen and Pemberton-8">Owen and Pemberton, p. 8.</ref> the explorers first encountered the animal by hearing its far-reaching vocalisations at night.<ref>Owen and Pemberton, p. 7.</ref> Related names that were used in the 19th century were ''Sarcophilus satanicus'' ("Satanic flesh-lover") and ''Diabolus ursinus'' ("bear devil"), all due to early misconceptions of the species as implacably vicious.<ref name="Owen and Pemberton-8"/> The Tasmanian devil (''Sarcophilus harrisii'') belongs to the family [[Dasyuridae]]. The genus ''Sarcophilus'' contains two other species, known only from [[Pleistocene]] fossils: ''S. laniarius'' and ''S. moomaensis''. [[Phylogenetics|Phylogenetic]] analysis shows that the Tasmanian devil is most closely related to [[quoll]]s.<ref name=Krajewski1992/> According to Pemberton, the possible ancestors of the devil may have needed to climb trees to acquire food, leading to a growth in size and the hopping gait of many marsupials. He speculated that these adaptations may have caused the contemporary devil's peculiar gait.<ref name="Owen and Pemberton-34">Owen and Pemberton, p. 34.</ref> The specific lineage of the Tasmanian devil is theorised to have emerged during the [[Miocene]], molecular evidence suggesting a split from the ancestors of quolls between 10 and 15 million years ago,<ref name=PwP16>{{cite book|author1=Krajewski, Carey |author2=Westerman, Michael |year=2003|title=Predators with Pouches: The Biology of Carnivorous Marsupials|chapter=Molecular Systematics of Dasyuromorpha|page=16|editor1=Jones, Menna |editor2=Dickman, Chris |editor3=Archer, Mike |publisher=CSIRO Publishing|location=Collingwood, Victoria|isbn=0-643-06634-9}}</ref> when severe [[climate change]] came to bear in Australia, transforming the climate from warm and moist to an arid, dry ice age, resulting in mass extinctions.<ref name="Owen and Pemberton-34"/> As most of their prey died of the cold, only a few carnivores survived, including the ancestors of the [[quoll]] and [[thylacine]]. It is speculated that the devil lineage may have arisen at this time to fill a niche in the ecosystem, as a scavenger that disposed of carrion left behind by the selective-eating thylacine.<ref name="Owen and Pemberton-34"/> The extinct ''[[Glaucodon]] ballaratensis'' of the [[Pliocene]] age has been dubbed an intermediate species between the quoll and devil.<ref>{{cite book|last1=Long|first1=John A.|author-link1=John A. Long |last2=Archer|first2=Michael|author-link2=Mike Archer (paleontologist) |last3=Flannery|first3=Timothy |last4=Hand|first4=Suzanne |title=Prehistoric Mammals of Australia and New Guinea: One Hundred Million Years of Evolution|url=https://books.google.com/books?id=92yhnRHdxSoC&pg=PA55|year=2002|publisher=University of New South Wales Press|isbn=978-0-8018-7223-5|page=55}}</ref> [[File:Jenolan Caves Tasmanian Devil jaw.jpg|thumb|right|A jawbone found in the mainland [[Jenolan Caves]]]] Fossil deposits in limestone caves at [[Naracoorte, South Australia]], dating to the [[Miocene]] include specimens of ''S. laniarius'', which were around 15% larger and 50% heavier than modern devils.<ref>Owen and Pemberton, p. 35.</ref> Older specimens believed to be 50–70,000 years old were found in [[Darling Downs]] in [[Queensland]] and in [[Western Australia]].<ref name="Owen and Pemberton-36"/> It is not clear whether the modern devil evolved from ''S. laniarius'', or whether they coexisted at the time.<ref name="Owen and Pemberton-36">Owen and Pemberton, p. 36.</ref> Richard Owen argued for the latter hypothesis in the 19th century, based on fossils found in 1877 in [[New South Wales]].<ref name="Owen and Pemberton-36"/> Large bones attributed to ''S. moornaensis'' have been found in New South Wales,<ref name="Owen and Pemberton-36"/> and it has been conjectured that these two extinct larger species may have hunted and scavenged.<ref name="Owen and Pemberton-36"/> It is known that there were several [[genus|genera]] of thylacine millions of years ago, and that they ranged in size, the smaller being more reliant on foraging.<ref>Owen and Pemberton, p. 37.</ref> As the devil and thylacine are similar, the extinction of the co-existing thylacine genera has been cited as evidence for an analogous history for the devils.<ref name="Owen and Pemberton-38">Owen and Pemberton, p. 38.</ref> It has been speculated that the smaller size of ''S. laniarius'' and ''S. moornaensis'' allowed them to adapt to the changing conditions more effectively and survive longer than the corresponding thylacines.<ref name="Owen and Pemberton-38"/> As the extinction of these two species came at a similar time to human habitation of Australia, hunting by humans and land clearance have been mooted as possible causes.<ref name="Owen and Pemberton-39">Owen and Pemberton, p. 39.</ref> Critics of this theory point out that as indigenous Australians only developed [[boomerang]]s and spears for hunting around 10,000 years ago, a critical fall in numbers due to systematic hunting is unlikely. They also point out that caves inhabited by Aborigines have a low proportion of bones and rock paintings of devils, and suggest that this is an indication that it was not a large part of indigenous lifestyle. A scientific report in 1910 claimed that Aborigines preferred the meat of herbivores rather than carnivores.<ref>Owen and Pemberton, pp. 40–42.</ref> The other main theory for the extinction was that it was due to the climate change brought on by the most recent ice age.<ref name="Owen and Pemberton-39"/> ===Genetics=== [[File:Karyotype of Tasmanian devil (Sarcophilus Harrisii).png|right|thumb|[[Karyotype]] of male Tasmanian devil]] The Tasmanian devil's [[genome]] was sequenced in 2010 by the [[Wellcome Trust Sanger Institute]].<ref>{{cite press release |url=http://www.sanger.ac.uk/news/view/2010-09-16-completed-genome-is-first-step-to-tackling-tasmanian-devil-facial-tumours |title=Completed genome is first step to tackling Tasmanian devil facial tumours|date=16 September 2010|access-date=10 December 2015|publisher=[[Wellcome Trust Sanger Institute]]}}</ref> Like all dasyurids, the devil has 14 chromosomes.<ref>Tyndale-Biscoe, p. 143.</ref> 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.<ref name=mj04/> According to a study by Menna Jones, "[[gene flow]] appears extensive up to {{cvt|50|km}}", meaning a high assignment rate to source or close neighbour populations "in agreement with movement data. At larger scales ({{cvt|150|–|250|km|disp=or}}), gene flow is reduced but there is no evidence for isolation by distance".<ref name=mj04/> [[Foster's rule|Island effects]] may also have contributed to their low genetic diversity. Periods of low population density may also have created moderate [[population bottleneck]]s, reducing genetic diversity.<ref name=mj04/> Low genetic diversity is thought to have been a feature in the Tasmanian devil population since the mid-[[Holocene]].<ref name="morris&austin2012"/> Outbreaks of [[devil facial tumour disease]] (DFTD) cause an increase in [[inbreeding]].<ref name=LachishMiller2010/> A sub-population of devils in the north-west of the state is genetically distinct from other devils,<ref name=fed>{{cite web|url=http://www.environment.gov.au/cgi-bin/sprat/public/publicspecies.pl?taxon_id=299|title=''Sarcophilus harrisii'' – Tasmanian Devil |work=Species Profile and Threats Database |publisher=Department of Sustainability, Environment, Water, Population and Communities|access-date=30 September 2010}}</ref> but there is some exchange between the two groups.<ref name=draft>{{cite web |year=2010 |url=http://dpipwe.tas.gov.au/Documents/Draft-Tasmanian-Devil-Recovery-Plan.pdf |title=Draft Recovery Plan for the Tasmanian devil (''Sarcophilus harrisii'')|publisher=Department of Primary Industries, Parks, Water and Environment |location=Hobart |access-date=3 September 2015}}</ref> One strand conformation polymorphism analysis (OSCP) on the [[major histocompatibility complex]] (MHC) [[MHC class I|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.<ref name=sid/> 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.<ref name=sid/><!-- cites both previous --> 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.<ref name=sid/> Recent research has suggested that the wild population of devils are rapidly evolving a resistance to DFTD.<ref>{{cite journal|last1=Epstein|first1=Brendan|last2=Jones|first2=Menna|last3=Hamede|first3=Rodrigo|last4=Hendricks|first4=Sarah|last5=McCallum|first5=Hamish|last6=Murchison|first6=Elizabeth P.|last7=Schönfeld|first7=Barbara|last8=Wiench|first8=Cody|last9=Hohenlohe|first9=Paul|last10=Storfer|first10=Andrew|title=Rapid evolutionary response to a transmissible cancer in Tasmanian devils|journal=[[Nature Communications]]|date=30 August 2016|volume=7|pages=12684|doi=10.1038/ncomms12684|pmid=27575253|pmc=5013612|bibcode=2016NatCo...712684E}}</ref> ==Description== [[File:Tasmanian Devil markings.jpg|right|thumb|Two devils, one without any white markings. Around 16% of wild devils have no markings.|alt=Two devils, sitting side by side, the one of left with a white stripe under its neck. They stand on a dirt patch. Stones can be seen in the background.]] [[File:Animaldentition sarcophilusharrisii.png|thumb|Dentition, as illustrated in Knight's ''Sketches in Natural History'']] The Tasmanian devil is the largest surviving carnivorous [[marsupial]]. It has a squat, thick build, with a large head and a tail which is about half its body length. Unusually for a marsupial, its forelegs are slightly longer than its hind legs, and devils can run up to {{cvt|13|km/h}} for short distances. The fur is usually black, often with irregular white patches on the chest and rump (although approximately 16% of wild devils do not have white patches).<ref name=Guiler1983/><ref name="Owen and Pemberton-4647">Owen and Pemberton, pp. 46–47.</ref> These markings suggest that the devil is most active at dawn and dusk, and they are thought to draw biting attacks toward less important areas of the body, as fighting between devils often leads to a concentration of scars in that region.<ref name="Owen and Pemberton-4647"/> Males are usually larger than females, having an average head and body length of {{cvt|652|mm}}, a {{cvt|258|mm}} tail and an average weight of {{cvt|8|kg}}. Females have an average head and body length of {{cvt|570|mm}}, a {{cvt|244|mm}} tail and an average weight of {{cvt|6|kg}},<ref name=Guiler1983>{{cite book|author=Guiler, E. R.|year=1983|chapter=Tasmanian Devil|editor=Strahan, R.|title=The Australian Museum Complete Book of Australian Mammals|pages=27–28|publisher=Angus & Robertson|isbn=0-207-14454-0}}</ref> although devils in western Tasmania tend to be smaller.<ref name="Owen and Pemberton-118">Owen and Pemberton, p. 118.</ref> Devils have five long toes on their forefeet, four pointing to the front and one coming out from the side, which gives the devil the ability to hold food. The hind feet have four toes, and the devils have non-retractable claws.<ref name=draft/> The stocky devils have a relatively low [[centre of mass]].<ref>Owen and Pemberton, p. 52.</ref> Devils are fully grown at two years of age,<ref name=fed/> and few devils live longer than five years in the wild.<ref name=Jones2008/> Possibly the longest-lived Tasmanian devil recorded was [[Coolah (Tasmanian Devil)|Coolah]], a male devil which lived in captivity for more than seven years.<ref>Owen and Pemberton, p. 140.</ref> Born in January 1997 at the [[Cincinnati Zoo and Botanical Garden|Cincinnati Zoo]], Coolah died in May 2004 at the [[Fort Wayne Children's Zoo]].<ref name=UPI2004>{{Cite news|title=Last Tasmanian devil not in Australia dies|work=United Press International|date=19 May 2004|url=http://www.upi.com/Odd_News/2004/05/19/Last-Tasmanian-devil-not-in-Australia-dies/UPI-43691084999988/|access-date=28 November 2013}}</ref> The devil stores body fat in its tail, and healthy devils have fat tails.<ref name=DPIWEweb1>{{cite web|title=Tasmanian devil – Frequently Asked Questions|url=https://nre.tas.gov.au/wildlife-management/save-the-tasmanian-devil-program/frequently-asked-questions/tasmanian-devils-faqs|publisher=Department of Natural Resources and Environment Tasmania|access-date=28 June 2022|archive-date=24 May 2022|archive-url=https://web.archive.org/web/20220524183500/https://nre.tas.gov.au/wildlife-management/save-the-tasmanian-devil-program/frequently-asked-questions/tasmanian-devils-faqs|url-status=dead}}</ref> The tail is largely non-prehensile and is important to its physiology, social behaviour and locomotion. It acts as a counterbalance to aid stability when the devil is moving quickly.<ref name="Owen and Pemberton-46">Owen and Pemberton, p. 46.</ref> An ano-genital scent gland at the base of its tail is used to mark the ground behind the animal with its strong, pungent scent.<ref name=Pemberton1993/> The male has external testes in a pouch-like structure formed by lateral ventrocrural folds of the abdomen, which partially hides and protects them. The testes are subovoid in shape and the mean dimensions of 30 testes of adult males was {{cvt|3.17|x|2.57|cm}}.<ref name="Guiler-64">Guiler (1970), p. 64.</ref> The female's pouch opens backwards, and is present throughout its life, unlike some other dasyurids.<ref name="Guiler-64"/> [[File:Tasmanian devil skeleton.jpg|thumb|left|Tasmanian devil skeleton on display at the [[Museum of Osteology]], [[Oklahoma City, Oklahoma]]]] The Tasmanian devil has the most powerful bite relative to body size of any living mammalian carnivore, with a [[Bite force quotient|Bite Force Quotient]] of 181 and exerting a canine bite force of {{cvt|553|N}}.<ref name=Wroe2005/><ref>{{Cite web|url=http://sydney.edu.au/news/84.html?newsstoryid=72|title=The Bite Club: comparative bite force in biting mammals|date=4 April 2005|publisher=The University of Sydney|language=en|archive-url=https://web.archive.org/web/20180118053123/http://sydney.edu.au/news/84.html?newsstoryid=72|archive-date=18 January 2018|access-date=2018-01-18}}</ref> The jaw can open to 75–80 degrees, allowing the devil to generate the large amount of power to tear meat and crush bones<ref name="Owen and Pemberton-46"/>—sufficient force to allow it to bite through thick metal wire.<ref>Owen and Pemberton, p. 20.</ref> The power of the [[jaw]]s is in part due to its comparatively large head. The teeth and jaws of Tasmanian devils resemble those of [[hyena]]s, an example of [[convergent evolution]].<ref>{{cite encyclopedia|encyclopedia=Encyclopædia Britannica|url=http://www.britannica.com/EBchecked/topic/583942/Tasmanian-devil|title=Tasmanian devil (marsupial) |access-date=16 March 2010}}</ref><ref name="Owen and Pemberton-64"/> Dasyurid teeth resemble those of primitive marsupials. Like all dasyurids, the devil has prominent canines and cheek teeth. It has three pairs of lower incisors and four pairs of upper incisors. These are located at the top of the front of the devil's mouth.<ref>Tyndale-Biscoe, pp. 142–143.</ref> Like dogs, it has 42 teeth, however, unlike dogs, its teeth are not replaced after birth but grow continuously throughout life at a slow rate.<ref name=DPIWEweb1/><ref name="Owen and Pemberton-64">Owen and Pemberton, p. 64.</ref> It has a "highly carnivorous [[dentition]] and [[wikt:trophic|trophic]] adaptations for bone consumption".<ref name="Owen and Pemberton-44"/> The devil has long claws that allow it to dig burrows and seek subterranean food easily and grip prey or mates strongly.<ref name="Owen and Pemberton-64"/> The teeth and claw strength allow the devil to attack wombats up to {{cvt|30|kg}} in weight.<ref name="Owen and Pemberton-44">Owen and Pemberton, p. 44.</ref> The large neck and forebody that give the devil its strength also cause this strength to be biased towards the front half of the body; the lopsided, awkward, shuffling gait of the devil is attributed to this.<ref>Owen and Pemberton, p. 53.</ref> The devil has long [[Vibrissae|whiskers]] on its face and in clumps on the top of the head. These help the devil locate prey when foraging in the dark, and aid in detecting when other devils are close during feeding.<ref name="Owen and Pemberton-64"/> The whiskers can extend from the tip of the chin to the rear of the jaw and can cover the span of its shoulder.<ref name="Owen and Pemberton-64"/> Hearing is its dominant sense, and it also has an excellent sense of smell, which has a range of {{cvt|1|km}}.<ref name=DPIWEweb1/><ref name="Owen and Pemberton-64"/> The devil, unlike other marsupials, has a "well-defined, saddle-shaped [[ectotympanic]]".<ref name=Wroe1999/> Since devils hunt at night, their vision seems to be strongest in [[monochromacy|black and white]]. In these conditions they can detect moving objects readily, but have difficulty seeing stationary objects.<ref name=DPIWEweb1/> ==Distribution and habitat== The Tasmanian devil was formerly present across mainland Australia, but became extinct there 3,500 years ago, co-incident with the extinction of the Thylacine from the region. A number of causal factors for the extinction have been proposed, including the introduction of the [[dingo]], intensification of human activity, as well as climatic change.<ref>{{Cite journal |last1=White |first1=Lauren C. |last2=Saltré |first2=Frédérik |last3=Bradshaw |first3=Corey J. A. |last4=Austin |first4=Jeremy J. |date=January 2018 |title=High-quality fossil dates support a synchronous, Late Holocene extinction of devils and thylacines in mainland Australia |journal=Biology Letters |language=en |volume=14 |issue=1 |pages=20170642 |doi=10.1098/rsbl.2017.0642 |issn=1744-9561 |pmc=5803592 |pmid=29343562}}</ref> Devils are found in all habitats on the island of Tasmania, including the outskirts of urban areas, and are distributed throughout the Tasmanian mainland and on [[Robbins Island (Tasmania)|Robbins Island]] (which is connected to mainland Tasmania at [[low tide]]).<ref name=vul/> The north-western population is located west of the [[Forth River, Australia|Forth River]] and as far south as [[Macquarie Harbour|Macquarie Heads]].<ref name="iucn" /> Previously, they were present on [[Bruny Island]] from the 19th century, but there have been no records of them after 1900.<ref name="iucn" /> They were illegally introduced to [[Badger Island]] in the mid-1990s but were removed by the Tasmanian government by 2007. Although the Badger Island population was free from DFTD, the removed individuals were returned to the Tasmanian mainland, some to infected areas.<ref>{{cite web |url=https://stors.tas.gov.au/download/1539284 |title=The Tasmanian Devil – Biology, Facial Tumour Disease and Conservation |first=Allen |last=Greer }}</ref> A study has modelled the reintroduction of DFTD-free Tasmanian devils to the Australian mainland in areas where [[dingo]]es are sparse. It is proposed that devils would have fewer impacts on both livestock and native fauna than dingoes, and that the mainland population could act as an additional [[insurance population]].<ref>{{cite news|first1=Daniel|last1=Hunter|first2=Mike|last2=Letnic|title=Bringing devils back to the mainland could help wildlife conservation|url=https://theconversation.com/bringing-devils-back-to-the-mainland-could-help-wildlife-conservation-43121|access-date=24 August 2015|work=[[The Conversation (website)|The Conversation]]}}</ref> In September 2015, 20 immunised captive-bred devils were released into [[Narawntapu National Park]], Tasmania.<ref>{{cite news|first1=Lucy|last1=Shannon |first2=Ros|last2=Lehman|title=Release of captive bred Tasmanian devils hailed as turning point in fight against disease|url=http://www.abc.net.au/news/2015-09-26/release-of-captive-bred-tasmania-devils-marks-milestone/6807058|access-date=26 September 2015|work=[[ABC News (Australia)|ABC News]]|date=26 September 2015}}</ref> Two later died from being hit by cars.<ref>{{cite news|last=Gramenz|first=Emilie|title=Two of 20 immunised Tasmanian devils released into wild killed on road days after release|url=http://www.abc.net.au/news/2015-09-30/tasmanian-devils-killed-on-road-days-after-being-released/6816966|access-date=30 September 2015|work=ABC News|date=30 September 2015}}</ref> The "core habitat" of the devils is considered to be within the "[[Climate of Tasmania|low to moderate annual rainfall zone]] of eastern and north-western Tasmania".<ref name=draft/> Tasmanian devils particularly like dry [[sclerophyll]] forests and coastal woodlands.<ref name=Fischer2001/> Although they are not found at the highest altitudes of Tasmania, and their [[population density]] is low in the [[Gymnoschoenus sphaerocephalus|button grass]] plains in the [[South West Wilderness|south-west of the state]], their population is high in dry or mixed sclerophyll forests and coastal heaths. Devils prefer open forest to tall forest, and dry rather than wet forests.<ref name=fed/> They are also found near roads where roadkill is prevalent, although the devils themselves are often killed by vehicles while retrieving the carrion.<ref name=vul/> According to the Threatened Species Scientific Committee, their versatility means that habitat modification from destruction is not seen as a major threat to the species.<ref name=vul/> The devil is directly linked to the ''[[Dasyurotaenia robusta]]'', a [[tapeworm]] which is classified as Rare under the Tasmanian Threatened Species Protection Act 1995. This tapeworm is found only in devils.<ref name=fed/> In late 2020, Tasmanian devils were reintroduced to mainland Australia in a sanctuary run by [[Aussie Ark]] in the [[Barrington Tops]] area of [[New South Wales]]. This was the first time devils had lived on the Australian mainland in over 3,000 years.<ref>{{Cite web|date=2020-10-05|title=Tasmanian devils return to mainland Australia for first time in 3,000 years|url=https://www.nationalgeographic.com/animals/2020/10/tasmanian-devils-return-to-mainland-australia/|archive-url=https://web.archive.org/web/20201006051259/https://www.nationalgeographic.com/animals/2020/10/tasmanian-devils-return-to-mainland-australia/|url-status=dead|archive-date=6 October 2020|access-date=2020-10-18|website=Animals|language=en}}</ref> 26 adult devils were released into the {{cvt|400|ha}} [[protected area]], and by late April 2021, seven [[joey (marsupial)|joeys]] had been born, with up to 20 expected by the end of the year.<ref>{{cite web | last=Conroy | first=Gemma | title=Tasmanian devils give birth in semi-wild sanctuary on the mainland | website=ABC News|publisher= [[Australian Broadcasting Corporation]] | date=27 May 2021 | url=https://www.abc.net.au/news/science/2021-05-27/tasmanian-devils-give-birth-in-semi-wild-sanctuary-on-mainland/100169686 | access-date=30 May 2021}}</ref> ==Ecology and behaviour== [[File:Tasmanian Devil resting.jpg|thumb|Although Tasmanian devils are nocturnal, they like to rest in the sun. [[Scar]]ring from fighting is visible next to this devil's left eye.|alt=A devil lying belly down on dry scrub grass and dead leaves. It has stretched its front legs out in front of its face.]] The Tasmanian devil is a [[keystone species]] in the ecosystem of Tasmania.<ref name="STTDNSept10">{{cite news|title=Hello ... or Goodbye|url=http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/DevilNews_Sep_2010.pdf|access-date=26 October 2010|newspaper=Save The Tasmanian Devil Newsletter|page=3|date=September 2010|archive-date=17 February 2011|archive-url=https://web.archive.org/web/20110217184444/http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/DevilNews_Sep_2010.pdf|url-status=dead}}</ref> It is a [[nocturnal animal|nocturnal]] and [[crepuscular]] hunter, spending the days in dense bush or in a hole.<ref name=Fischer2001/> It has been speculated that nocturnalism may have been adopted to avoid predation by eagles and humans.<ref name="Owen and Pemberton-129"/> Young devils are predominantly crepuscular.<ref name="Owen and Pemberton-69"/> There is no evidence of [[torpor]].<ref name=jb98/> Young devils can climb trees, but this becomes more difficult as they grow larger.<ref name=ors/><ref>{{cite news|title=Young devil displays gnarly climbing technique|url=http://www.tassiedevil.com.au/tasdevil.nsf/news/E88F860D80C1554BCA2578780012A8E3|access-date=4 May 2011|newspaper=Save The Tasmanian Devil Program|date=3 May 2011|archive-date=30 June 2017|archive-url=https://web.archive.org/web/20170630062135/http://www.tassiedevil.com.au/tasdevil.nsf/news/E88F860D80C1554BCA2578780012A8E3|url-status=dead}}</ref> Devils can scale trees of trunk diameter larger than {{cvt|40|cm}}, which tend to have no small side branches to hang onto, up to a height of around {{cvt|2.5|–|3|m}}. Devils that are yet to reach maturity can climb shrubs to a height of {{cvt|4|m}}, and can climb a tree to {{cvt|7|m}} if it is not vertical.<ref name=jb00/> Adult devils may eat young devils if they are very hungry, so this climbing behaviour may be an adaptation to allow young devils to escape.<ref>{{cite web|url=http://dpipwe.tas.gov.au/wildlife-management/animals-of-tasmania/mammals/carnivorous-marsupials-and-bandicoots/tasmanian-devil/tasmanian-devil-information-for-kids/tasmanian-devil-facts-for-kids|publisher=Department of Primary Industries, Parks, Water and Environment|title=Tasmanian Devil Facts for Kids|date=1 September 2014|access-date=3 September 2015|url-status=dead|archive-url=https://web.archive.org/web/20150825024852/http://dpipwe.tas.gov.au/wildlife-management/animals-of-tasmania/mammals/carnivorous-marsupials-and-bandicoots/tasmanian-devil/tasmanian-devil-information-for-kids/tasmanian-devil-facts-for-kids|archive-date=25 August 2015|df=dmy-all}}</ref> Devils can also swim and have been observed crossing rivers that are {{cvt|50|m}} in width, including icy cold waterways, apparently enthusiastically.<ref name=ors>Owen and Pemberton, pp. 21–22.</ref> Tasmanian devils do not form packs, but rather spend most of their time alone once weaned.<ref name=Fischer2001/><ref name="Owen and Pemberton-69"/> Classically considered as [[solitary animals]], their social interactions were poorly understood. However, a [[field study]] published in 2009 shed some light on this. Tasmanian devils in [[Narawntapu National Park]] were fitted with proximity sensing [[Tracking collar|radio collars]] which recorded their interactions with other devils over several months from February to June 2006. This revealed that all devils were part of a single huge contact network, characterised by male-female interactions during mating season, while female–female interactions were the most common at other times, although frequency and patterns of contact did not vary markedly between seasons. Previously thought to fight over food, males only rarely interacted with other males.<ref name=Hamede/><!-- ref cites previous 6 sentences --> Hence, all devils in a region are part of a single social network.<ref>{{Cite news|title=Social Networking Study Reveals Threat To Tasmanian Devils|url=https://www.sciencedaily.com/releases/2009/08/090819064033.htm|access-date=26 August 2010|newspaper=Science Daily|date=19 August 2009}}</ref> They are considered to be [[Territory (animal)|non-territorial]] in general, but females are territorial around their dens.<ref name=DPIWEweb1/> This allows a higher total mass of devils to occupy a given area than territorial animals, without conflict.<ref name=oran/> Tasmanian devils instead occupy a [[home range]].<ref>{{cite web |url=http://dpipwe.tas.gov.au/wildlife-management/animals-of-tasmania/mammals/carnivorous-marsupials-and-bandicoots/tasmanian-devil |access-date=4 September 2015 |title=Tasmanian Devil |date=13 November 2014 |publisher=Department of Primary Industries, Parks, Water and Environment |url-status=dead |archive-url=https://web.archive.org/web/20150908100030/http://dpipwe.tas.gov.au/wildlife-management/animals-of-tasmania/mammals/carnivorous-marsupials-and-bandicoots/tasmanian-devil |archive-date=8 September 2015 |df=dmy-all }}</ref> In a period of between two and four weeks, devils' home ranges are estimated to vary between {{cvt|4|and|27|km2}}, with an average of {{cvt|13|km2}}.<ref name=fed/> The location and geometry of these areas depend on the distribution of food, particularly [[Wallaby|wallabies]] and [[pademelon]]s nearby.<ref name=oran/> Devils use three or four dens regularly. Dens formerly owned by [[wombats]] are especially prized as maternity dens because of their security. Dense vegetation near creeks, thick grass tussocks, and caves are also used as dens. Adult devils use the same dens for life. It is believed that, as a secure den is highly prized, some may have been used for several centuries by generations of animals.<ref name=oran/> Studies have suggested that food security is less important than den security, as [[habitat destruction]] that affects the latter has had more effect on mortality rates.<ref name=oran/> Young pups remain in one den with their mother, and other devils are mobile,<ref name=oran>Owen and Pemberton, pp. 76–77.</ref> changing dens every 1–3 days and travelling a [[mean]] distance of {{cvt|8.6|km}} every night.<ref>{{cite web|url=http://www.environment.gov.au/biodiversity/threatened/species/pubs/sarcophilus-harrisii-listing.pdf|publisher=Department of Sustainability, Environment, Water, Population and Communities|title=Advice to the Minister for the Environment and Heritage from the Threatened Species Scientific Committee (the Committee) on Amendments to the list of Threatened Species under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act)|access-date=26 October 2010|author=Threatened Species Scientific Committee (the Committee) on Amendments to the list of Threatened Species}}</ref> However, there are also reports that an upper bound can be {{cvt|50|km}} per night. They choose to travel through lowlands, saddles and along the banks of creeks, particularly preferring carved-out tracks and livestock paths and eschewing steep slopes and rocky terrain.<ref name=fed/><ref name="Owen and Pemberton-118"/> The amount of movement is believed to be similar throughout the year, except for mothers who have given birth recently.<ref name=fed/> The similarity in travel distances for males and females is unusual for sexually dimorphic, solitary carnivores. As a male needs more food, he will spend more time eating than travelling. Devils typically make circuits of their home range during their hunts.<ref name="oran"/> In areas near human habitation, they are known to steal clothes, blankets and pillows and take them for use in dens in wooden buildings.<ref>Owen and Pemberton, pp. 23–24.</ref> While the [[Dasyuridae|dasyurids]] have similar diet and anatomy, differing body sizes affect [[thermoregulation]] and thus behaviour.<ref>Tyndale-Biscoe, p. 148.</ref> In ambient temperatures between {{cvt|5|and|30|°C}}, the devil was able to maintain a body temperature between {{cvt|37.4|and|38|°C}}. When the temperature was raised to {{cvt|40|°C}}, and the humidity to 50%, the devil's body temperature spiked upwards by {{cvt|2|°C-change}} within 60 minutes, but then steadily decreased back to the starting temperature after a further two hours, and remained there for two more hours. During this time, the devil drank water and showed no visible signs of discomfort, leading scientists to believe that sweating and evaporative cooling is its primary means of heat dissipation.<ref>Tyndale-Biscoe, pp. 147–149.</ref> A later study found that devils pant but do not sweat to release heat.<ref name=draft/> In contrast, many other marsupials were unable to keep their body temperatures down.<ref name="Tyndale-149">Tyndale-Biscoe, p. 149.</ref> As the smaller animals have to live in hotter and more arid conditions to which they are less well-adapted, they take up a nocturnal lifestyle and drop their body temperatures during the day, whereas the devil is active in the day and its body temperature varies by {{cvt|1.8|°C-change}} from its minimum at night to the maximum in the middle of the day.<ref>Tyndale-Biscoe, pp. 148–149.</ref> The standard [[metabolic rate]] of a Tasmanian devil is 141 kJ/kg (15.3 [[Calorie|kcal]]/lb) per day, many times lower than smaller marsupials. A {{cvt|5|kg}} devil uses {{cvt|712|kJ}} per day. The field metabolic rate is 407 kJ/kg (44.1 kcal/lb).<ref name="Tyndale-149"/> Along with quolls, Tasmanian devils have a metabolic rate comparable to non-carnivorous marsupials of a similar size. This differs from placental carnivores, which have comparatively high basal metabolic rates.<ref name=Cooper2010/> A study of devils showed a loss of weight from {{cvt|7.9|to|7.1|kg}} from summer to winter, but in the same time, daily energy consumption increased from {{cvt|2591|to|2890|kJ}}. This is equivalent to an increase in food consumption from {{cvt|518|to|578|g}}.<ref name="Tyndale-150">Tyndale-Biscoe, p. 150.</ref> The diet is protein-based with 70% water content. For every {{cvt|1|g}} of insects consumed, {{cvt|3.5|kJ}} of energy are produced, while a corresponding amount of wallaby meat generated {{cvt|5.0|kJ}}.<ref name="Tyndale-150"/> In terms of its body mass, the devil eats only a quarter of the [[eastern quoll]]'s intake,<ref name="Tyndale-150"/> allowing it to survive longer during food shortages. ===Feeding=== [[File:Devil-eating-roadkill.jpg|right|thumb|A devil eating roadkill|alt=A black devil standing on a patch of cut grass in a paddock, next to a wire fence. It is biting into the torn carcass of an animal.]] 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 [[wombat]]s because of the ease of predation and high fat content, it will eat all small native mammals such as [[wallabies]],<ref name="ADW">{{cite web |url=https://animaldiversity.org/accounts/Sarcophilus_harrisii/ |title=Sarcophilus harrisii (Tasmanian devil) |website=animaldiversity.org}}</ref> [[bettong]] and [[potoroo]]s, domestic mammals (including sheep and rabbits),<ref name="ADW"/> birds (including [[penguin]]s),<ref>{{Cite web|title=Tasmanian devils on tiny Australian island wipe out thousands of penguins|url=https://www.9news.com.au/national/tasmanian-devils-on-tiny-australian-island-wipe-out-thousands-of-penguins/9a6da8f3-d6df-44b9-bd63-53f818858125|access-date=2021-06-22|website=www.9news.com.au|date=22 June 2021 }}</ref> fish, fruit, vegetable matter, insects, tadpoles, frogs and reptiles. Their diet is widely varied and depends on the food available.<ref name=DPIWEweb1/><ref name=of>Owen and Pemberton, pp. 11–13.</ref><ref name=orit/><ref>Owen and Pemberton, p. 108.</ref> 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.<ref name="Owen and Pemberton-44"/> 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,<ref name=of/> and eat the legs of otherwise robust sheep that have slipped in wooden shearing sheds, leaving their legs dangling below.<ref name=ors/> Other unusual matter observed in devil scats includes collars and tags of devoured animals, intact echidna spines, pencil, plastic and jeans.<ref name="Owen and Pemberton-129">Owen and Pemberton, p. 129.</ref> Devils can bite through metal traps, and tend to reserve their strong jaws for escaping captivity rather than breaking into food storage.<ref name="Owen and Pemberton-129"/> Due to their relative lack of speed, they cannot run down a wallaby or a rabbit, but they can attack animals that have become slow due to illness.<ref name=of/> They survey flocks of sheep by sniffing them from {{cvt|10|-|15|m}} away and attack if the prey is ill. The sheep stamp their feet in a show of strength.<ref name="Owen and Pemberton-129"/> Despite their lack of extreme speed, there have been reports that devils can run at {{cvt|25|km/h}} for {{cvt|1.5|km}}, 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.<ref name=ors/> Pemberton has reported that they can average {{cvt|10|km/h}} 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.<ref name=ors/> Devils can dig to forage [[corpse]]s, 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.<ref name=of/> 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.<ref name=Pemberton1993/> 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.<ref name=orit/> Tasmanian devils can eliminate all traces of a carcass of a smaller animal, devouring the bones and fur if desired.<ref>Owen and Pemberton, pp. 11–15, 20, 36.</ref> 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.<ref name="Owen and Pemberton-14">Owen and Pemberton, p. 14.</ref> 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.<ref name=of/> 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.<ref name=jb98/> Juvenile devils are sometimes known to climb trees;<ref>Owen and Pemberton, pp. 49–50.</ref> in addition to small vertebrates and invertebrates, juveniles climb trees to eat grubs and birds' eggs.<ref name="Owen and Pemberton-69"/> Juveniles have also been observed climbing into nests and capturing birds.<ref name=jb00/> 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.<ref name=jb00/> 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.<ref name=jb00/> [[File:Tasmanian Devils feeding.jpg|right|thumb|Three Tasmanian devils feeding. Eating is a social event for the Tasmanian devil, and groups of 2 to 5 are common.|alt=Three Tasmanian devils standing on bark chips huddled with their heads close together.]] Although they hunt alone,<ref name=DPIWEweb1/> there have been unsubstantiated claims of communal hunting, where one devil drives prey out of its habitat and an accomplice attacks.<ref name=of/> Eating is a social event for the Tasmanian devil. This combination of a solitary animal that eats communally makes the devil unique among carnivores.<ref name="Owen and Pemberton-69"/> Much of the noise attributed to the animal is a result of raucous communal eating, at which up to 12 individuals can gather,<ref name=Pemberton1993/> although groups of two to five are common;<ref>Owen and Pemberton, p. 71.</ref> 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.<ref name=of/> The amount of noise is correlated to the size of the carcass.<ref name=of/> The devils eat in accordance with a system. Juveniles are active at dusk, so they tend to reach the source before the adults.<ref name=orit>Owen and Pemberton, pp. 70–73.</ref> Typically, the [[dominance (ethology)|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.<ref name=orit/> When [[quoll]]s are eating a carcass, devils will tend to chase them away.<ref name=jb00/> This is a substantial problem for [[spotted-tailed quoll]]s, as they kill relatively large possums and cannot finish their meal before devils arrive. In contrast, the smaller [[eastern quoll]]s prey on much smaller victims, and can complete feeding before devils turn up.<ref name=jb00/> This is seen as a possible reason for the relatively small population of spotted-tailed quolls.<ref name=jb00/> A study of feeding devils identified twenty physical postures, including their characteristic vicious yawn, and eleven different vocal sounds, including clicks, shrieks and various types of [[growling|growls]], that devils use to communicate as they feed.<ref name=Pemberton1993/> They usually establish dominance by sound and physical posturing,<ref name=dinner>{{cite web |title=Devils at dinner |url=http://www.tassiedevil.com.au/tasdevil.nsf/Tasmaniandevils/EB75209E057BE43BCA2577BA0012D7AB?OpenDocument |publisher=Save the Tasmanian Devil |date=10 December 2010 |access-date=4 May 2011 |archive-date=20 March 2018 |archive-url=https://web.archive.org/web/20180320123807/http://www.tassiedevil.com.au/tasdevil.nsf/Tasmaniandevils/EB75209E057BE43BCA2577BA0012D7AB?OpenDocument |url-status=dead }}</ref> although fighting does occur.<ref name=Pemberton1993/> The white patches on the devil are visible to the night-vision of its colleagues.<ref name=orit/> Chemical gestures are also used.<ref name=orit/> Adult males are the most aggressive,<ref>Guiler (1992), pp. 8–10.</ref> and scarring is common.<ref>Owen and Pemberton, pp. 71–73.</ref> They can also stand on their hind legs and push each other's shoulders with their front legs and heads, similar to [[sumo wrestling]].<ref name=orit/> 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.<ref name=orit/> 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.<ref>Tyndale-Biscoe, p. 147.</ref> Devils are known to return to the same places to defecate, and to do so at a communal location, called a ''devil latrine''.<ref name="Owen and Pemberton-25">Owen and Pemberton, p. 25.</ref> It is believed that the communal defecation may be a means of communication that is not well understood.<ref name="Owen and Pemberton-25"/> Devil scats are very large compared to body size; they are on average {{cvt|15|cm}} long, but there have been samples that are {{cvt|25|cm}} in length.<ref name="Owen and Pemberton-25"/> They are characteristically grey in colour due to digested bones, or have bone fragments included.<ref name=fed/> 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.<ref>Owen and Pemberton, pp. 43–47.</ref> [[Wedge-tailed eagle]]s have a similar carrion-based diet to the devils and are regarded as competitors.<ref>Owen and Pemberton, pp. 60–62.</ref> 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.<ref>Owen and Pemberton, pp. 56–58.</ref> Both the Tasmanian devil and the quolls appears to have evolved up to 50 times faster than the average evolutionary rate amongst mammals.<ref>{{cite web|url=http://planetearth.nerc.ac.uk/news/story.aspx?id=1082|title=Mammals evolved at a steady pace|date=2011-10-26|archive-date=2011-12-30|archive-url=https://web.archive.org/web/20111230005839/http://planetearth.nerc.ac.uk/news/story.aspx?id=1082|access-date=14 May 2016}}</ref> ===Reproduction=== [[File:Tasmanina Devil development.PNG|thumb|500px|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.|alt=Black and white diagram indicating various points in a devil's development. Fur is absent until its development from days 50 to 85. The mouth begins round and the lips form from around days 20 to 25. The lips open from around days 80 to 85. They become unattached from their mother at around 100 to 105 days. The east are absent at birth and develop from around days 15 to 18. They are applied to the header until becoming erect from around 72 to 77 days. The eye slits develop around 20 days, and eyelashes between 50 and 55 days. The eyes open between 90 and 100 days.]] Females start to breed when they reach sexual maturity, typically in their second year. At this point, they become [[Estrus cycle|fertile]] once a year, producing multiple [[ovum|ova]] while in heat.<ref name=Guiler1970/> 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.<ref>Tyndale-Biscoe, p. 152.</ref> 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 [[mating|mate]] with the dominant male.<ref name=DPIWEweb1/><ref name=omate>Owen and Pemberton, pp. 64–66.</ref> 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.<ref name=omate/> Devils are not [[monogamy in animals|monogamous]], and females will mate with several males if not guarded after mating; males also reproduce with several females during a season.<ref name=DPIWEweb1/><ref name=omate/> Females have been shown to be selective in an attempt to ensure the best genetic offspring,<ref name=omate/> for example, fighting off the advances of smaller males.<ref name=draft/> Males often keep their mates in custody in the den, or take them along if they need to drink, lest they engage in infidelity.<ref name=omate/> Males can produce up to 16 offspring over their lifetime, while females average four mating seasons and 12 offspring.<ref name=omate/> Theoretically this means that a devil population can double on an annual basis and make the species insulated against high mortality.<ref>Owen and Pemberton, p. 66.</ref> The pregnancy rate is high; 80% of two-year-old females were observed with newborns in their pouches during the mating season.<ref name=omate/> More recent studies of breeding place the mating season between February and June, as opposed to between February and March.<ref name=fed/> [[Gestation]] lasts 21 days, and devils give birth to 20–30 young standing up,<ref name=DPIWEweb1/><ref name=omate/> each weighing approximately {{cvt|0.18|–|0.24|g}}.<ref name=Fischer2001/> [[Embryonic diapause]] does not occur.<ref name=Guiler1970/> 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 ({{cvt|0.26|–|0.43|cm|disp=or}}) than the rear limb ({{cvt|0.20|–|0.28|cm|disp=or}}), 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.<ref name=Guiler1970/> Tasmanian devil young are variously called "pups",<ref name=DPIWEweb1/> "joeys",<ref>{{cite news|title=Save The Tasmanian Devil Newsletter|url=http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/Devilnews_March_2010.pdf|access-date=2 October 2010|newspaper=Save The Tasmanian Devil Newsletter|date=March 2010|archive-date=17 February 2011|archive-url=https://web.archive.org/web/20110217184611/http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/Devilnews_March_2010.pdf|url-status=dead}}</ref> or "imps".<ref>{{cite web|title=Life Cycle of the Tasmanian Devil|url=http://www.tassiedevil.com.au/tasdevil.nsf/file/DD87BEF28EFEF6C3CA2576D20004BB2C/$file/STDP_Life%20Cycle%20Poster_A3.pdf|publisher=Save The Tasmanian Devil Program|access-date=2 October 2010|archive-date=17 February 2011|archive-url=https://web.archive.org/web/20110217174526/http://www.tassiedevil.com.au/tasdevil.nsf/file/DD87BEF28EFEF6C3CA2576D20004BB2C/$file/STDP_Life%20Cycle%20Poster_A3.pdf|url-status=dead}}</ref> 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.<ref name=DPIWEweb1/><ref name=omate/> On average, more females survive than males,<ref name=Guiler1970/> and up to 60% of young do not survive to maturity.<ref name="Owen and Pemberton-69">Owen and Pemberton, p. 69.</ref> 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.<ref name=Chuang2013/> Inside the pouch, the nourished young develop quickly. In the second week, the [[rhinarium]] becomes distinctive and heavily pigmented.<ref name=Guiler1970/> At 15 days, the [[pinna (anatomy)|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 {{cvt|1|cm}} long, and by the time the ear becomes erect, it is between {{cvt|1.2|and|1.6|cm}}. Eyelids are apparent at 16 days, whiskers at 17 days, and the lips at 20 days.<ref name=Guiler1970/> The devils can make squeaking noises after eight weeks, and after around 10–11 weeks, the lips can open.<ref name=Guiler1970/> Despite the formation of eyelids, they do not open for three months, although eyelashes form at around 50 days.<ref name=Guiler1970/> 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.<ref name=Guiler1970/> [[File:Young tasmanian devils.jpg|right|Three young devils sunbathing|thumb|alt=A view from directly above three devils lying with bodies almost touching, on dry leaves, dirt and rocks, under bright sunshine.]] The devils have a complete set of facial [[vibrissae]] and ulnar carpels, although it is devoid of [[wikt:anconeal|anconeal]] vibrissae. During the third week, the [[wikt:mystacial|mystacials]] and ulnarcarpals are the first to form. Subsequently, the [[wikt:infraorbital|infraorbital]], interramal, [[wikt:supraorbital|supraorbital]] and submental vibrissae form. The last four typically occur between the 26th and 39th day.<ref name=Guiler1970/> 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.<ref name=Guiler1970/> They leave the pouch 105 days after birth, appearing as small copies of the parent and weighing around {{cvt|200|g}}.<ref name=Guiler1970/> Zoologist Eric Guiler recorded its size at this time as follows: a [[crown-snout length]] of {{cvt|5.87|cm}}, tail length of {{cvt|5.78|cm}}, [[Pes (anatomy)|pes]] length {{cvt|2.94|cm}}, [[Manus (anatomy)|manus]] {{cvt|2.30|cm}}, shank {{cvt|4.16|cm}}, forearm {{cvt|4.34|cm}} and [[crown-rump length]] is {{cvt|11.9|cm}}.<ref name=Guiler1970/> During this period, the devils lengthen at a roughly linear rate.<ref name=Guiler1970/> 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.<ref name=Guiler1970/><ref>Guiler (1992), pp. 16–22.</ref> The milk contains a higher amount of iron than the milk of placental mammals.<ref name=draft/> In Guiler's 1970 study, no females died while rearing their offspring in the pouch. After leaving the pouch, the devils grow by around {{cvt|0.5|kg}} a month until they are six months old.<ref name=Guiler1970/> While most pups will survive to be weaned,<ref name=fed/> Guiler reported that up to three fifths of devils do not reach maturity.<ref name="Owen and Pemberton-69"/> As juveniles are more crepuscular than adults, their appearance in the open during summer gives the impression to humans of a population boom.<ref name="Owen and Pemberton-69"/> 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.<ref name=Sinn2014/> ==Conservation status== [[File:Didelphis cynocephala and Didelphis ursina, 1808.jpg|right|thumb|An 1808 impression featuring the Tasmanian devil and a [[thylacine]] by [[George Prideaux Robert Harris|George Harris]]|alt=A black and white drawing of a devil, which is in the upper half of the picture, facing right, and a thylacine in the lower half, facing left. Both are shown in profile and depicted on a matting of grass or other vegetation.]] The cause of the devil's disappearance from the mainland is unclear, but their decline seems to coincide with an abrupt change in climate and the expansion across the mainland of [[indigenous Australian]]s and [[dingo]]es.<ref>{{Cite journal|last1=Johnson|first1=C. N.|last2=Wroe|first2=S.|date=2016-07-27|title=Causes of extinction of vertebrates during the Holocene of mainland Australia: arrival of the dingo, or human impact?|url=https://journals.sagepub.com/doi/10.1191/0959683603hl682fa|journal=The Holocene|volume=13|issue=6|pages=941–948|language=en|doi=10.1191/0959683603hl682fa|bibcode=2003Holoc..13..941J |s2cid=15386196}}</ref><ref>{{Cite journal|last1=Prowse|first1=Thomas A. A.|last2=Johnson|first2=Christopher N.|last3=Bradshaw|first3=Corey J. A.|last4=Brook|first4=Barry W.|date=2014|title=An ecological regime shift resulting from disrupted predator–prey interactions in Holocene Australia|url=https://esajournals.onlinelibrary.wiley.com/doi/abs/10.1890/13-0746.1|journal=Ecology|language=en|volume=95|issue=3|pages=693–702|doi=10.1890/13-0746.1|pmid=24804453|bibcode=2014Ecol...95..693P |issn=1939-9170}}</ref> However, whether it was direct hunting by people, competition with dingoes, changes brought about by the increasing human population, who by 3000 years ago were using all habitat types across the continent, or a combination of all three, is unknown; devils had coexisted with dingoes on the mainland for around 3000 years.<ref name=Johnson2003/><!--cites previous 2 or 3 sentences--> Brown has also proposed that the [[El Niño–Southern Oscillation]] (ENSO) grew stronger during the Holocene, and that the devil, as a scavenger with a short life span, was highly sensitive to this.<ref name=Brown2006/> In dingo-free Tasmania,<ref>Owen and Pemberton, p. 40.</ref> carnivorous marsupials were still active when Europeans arrived. The extermination of the [[thylacine]] after the arrival of the Europeans is well known,<ref>Owen and Pemberton, p. 43.</ref> but the Tasmanian devil was threatened as well.<ref name="PWS" /> Habitat disruption can expose dens where mothers raise their young. This increases mortality, as the mother leaves the disturbed den with her pups clinging to her back, making them more vulnerable.<ref>Owen and Pemberton, pp. 75–76.</ref> Cancer in general is a common cause of death in devils.<ref>Owen and Pemberton, p. 171.</ref> In 2008, high levels of potentially [[carcinogenic]] flame retardant chemicals were found in Tasmanian devils. Preliminary results of tests ordered by the Tasmanian government on chemicals found in fat tissue from 16 devils have revealed high levels of [[hexabromobiphenyl]] (BB153) and "reasonably high" levels of [[decabromodiphenyl ether]] (BDE209).<ref>{{cite news|last=Denholm |first=M. |url=http://www.news.com.au/story/0,23599,23087523-421,00.html |title=Cancer agents found in Tasmanian devils |publisher=News Limited |date=22 January 2008 |access-date=30 September 2010 |url-status=dead |archive-url=https://web.archive.org/web/20090413035950/http://www.news.com.au/story/0%2C23599%2C23087523-421%2C00.html |archive-date=13 April 2009 }}</ref> The Save the Tasmanian Devil Appeal is the official fundraising entity for the Save the Tasmanian Devil Program. The priority is to ensure the survival of the Tasmanian devil in the wild. ===Population declines=== At least two major population declines, possibly due to disease epidemics, have occurred in recorded history: in 1909 and 1950.<ref name=Guiler1983/> The devil was also reported as scarce in the 1850s.<ref name=Bradshaw2005/> It is difficult to estimate the size of the devil population.<ref name="distimpactdftd"/> In the mid-1990s, the population was estimated at 130,000–150,000 animals,<ref name=fed/> but this is likely to have been an overestimate.<ref name="distimpactdftd"/> 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.<ref name=DPIWEweb1/> 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.<ref>{{cite web|last=Connellan|first=I|title=Tasmanian devils: Devil coast|url=http://www.australiangeographic.com.au/journal/tasmanian-devils-devil-coast.htm|work=Australian Geographic|access-date=22 August 2010|date=October–December 2008|archive-url=https://web.archive.org/web/20100830032310/http://www.australiangeographic.com.au/journal/tasmanian-devils-devil-coast.htm|archive-date=30 August 2010|url-status=dead|df=dmy-all}}</ref> The species was listed as vulnerable under the Tasmanian ''[[Threatened Species Protection Act 1995]]'' in 2005<ref>{{cite web|publisher=Department of the Environment and Heritage|date=July 2006|url=http://www.environment.gov.au/system/files/resources/45200a21-6135-4464-a5f6-1f9ac5ab9e7b/files/tasmanian-devil-policy.pdf|title=EPBC Policy Statement 3.6 – Tasmanian Devil (Sarcophilus harrisii)|access-date=3 September 2015}}</ref> and the Australian ''[[Environment Protection and Biodiversity Conservation Act 1999]]''<ref name=fed/> in 2006, which means that it is at risk of extinction in the "medium term".<ref name=vul>{{cite web|title=Advice to the Minister for the Environment, Heritage and the Arts from the Threatened Species Scientific Committee (the Committee) on Amendment to the list of Threatened Species under the Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act) Sarcophilus harrisii (Tasmanian Devil) Listing Advice|author=Beeton, Robert J. S.|publisher=Threatened Species Scientific Committee|url=http://www.environment.gov.au/biodiversity/threatened/species/pubs/299-listing-advice.pdf|access-date=23 October 2010|year=2009}}</ref> The [[IUCN]] classified the Tasmanian devil in the lower risk/least concern category in 1996, but in 2009 they reclassified it as endangered.<ref name="iucn" /> Appropriate wildlife refuges such as [[Savage River National Park]] in North West Tasmania provide hope for their survival. ===Culling=== The first European Tasmanian settlers ate Tasmanian devil, which they described as tasting like [[veal]].<ref name=Harris>{{cite book|author=Harris, G. P.|year=1807|chapter=Description of two species of Didelphis for Van Diemen's Land|title=Transactions of the Linnean Society of London|volume=IX}}</ref> 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.<ref name="Owen and Pemberton-9">Owen and Pemberton, p. 9.</ref> However, Guiler's research contended that the real cause of livestock losses was poor land management policies and feral dogs.<ref name="Owen and Pemberton-9"/> In areas where the devil is now absent, poultry has continued to be killed by [[quoll]]s. 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.<ref>Owen and Pemberton, p. 19.</ref> Over the next 100 years, trapping and poisoning<ref>Owen and Pemberton, pp. 19, 26–27.</ref> brought them to the brink of extinction.<ref name=PWS>{{cite web|url=http://www.parks.tas.gov.au/index.aspx?base=387|title=Tasmanian Devil, Sarcophilus harrisii|publisher=Parks & Wildlife Service Tasmania|access-date=26 September 2010|archive-url=https://web.archive.org/web/20110206221530/http://www.parks.tas.gov.au/index.aspx?base=387|archive-date=6 February 2011|url-status=dead}}</ref> After the death of the last thylacine in 1936,<ref>Paddle, p. 195.</ref> the Tasmanian devil was protected by law in June 1941 and the population slowly recovered.<ref name=PWS/> 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.<ref name="Owen and Pemberton-99">Owen and Pemberton, p. 99.</ref> 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.<ref>Owen and Pemberton, pp. 101–109.</ref> 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.<ref>Owen and Pemberton, pp. 118–119.</ref> Another report of overpopulation and livestock damage was reported in 1987.<ref>Owen and Pemberton, pp. 120–121.</ref> 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.<ref>Owen and Pemberton, pp. 127–129.</ref> 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.<ref name=vul/> Approximately 10,000 devils were killed per year in the mid-1990s.<ref name=draft/> 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.<ref name=LachishMcCallum/> 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.<ref name=BeetonMcCallum/> ===Road mortality=== [[File:Tasmanian Devil roadsign.jpg|right|thumb|A road sign telling drivers that there may be devils nearby|alt=A square-shaped metal tilted at 45 degrees on a metal post. The sign is painted yellow with a picture of a black devil in profile. It is at the side of a straight road cutting through wooded forest and two vehicles can be seen.]] Motor vehicles are a threat to localised populations of non-abundant Tasmanian mammals,<ref name="Hobday-2010"/><ref name="Jones, Menna E.-2000"/> 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.<ref name="Hobday-2010"/> 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.<ref name="Jones, Menna E.-2000"/> The vast majority of deaths occurred in the sealed portion of the road, believed to be due to an increase in speeds.<ref name="Jones, Menna E.-2000"/> 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.<ref name="Jones, Menna E.-2000"/> 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.<ref name="Owen and Pemberton-14"/> It was estimated that 3,392 devils, or 3.8–5.7% of the population, were being killed annually by vehicles in 2001–2004.<ref name=vul/> 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.<ref>{{cite web |title=Roadkill Project |publisher=Save the Tasmanian Devil |url=http://www.tassiedevil.com.au/tasdevil.nsf/Roadkill-Project/1A27B0F89FF95EF8CA2576D20077F70B |access-date=10 December 2015 |archive-date=18 March 2018 |archive-url=https://web.archive.org/web/20180318093646/http://www.tassiedevil.com.au/tasdevil.nsf/Roadkill-Project/1A27B0F89FF95EF8CA2576D20077F70B |url-status=dead }}</ref> On 25 September 2015, 20 immunised devils were microchipped and released in Narawntapu National Park. By 5 October four 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.<ref>{{cite web |url=http://www.tassiedevil.com.au/tasdevil.nsf/news/367C04BF953C460BCA257ED50002D816 |title=Devil deaths spark renewed plea for drivers to slow down |date=5 October 2015 |publisher=Save the Tasmanian Devil |access-date=10 December 2015 |archive-date=30 June 2017 |archive-url=https://web.archive.org/web/20170630054955/http://www.tassiedevil.com.au/tasdevil.nsf/news/367C04BF953C460BCA257ED50002D816 |url-status=dead }}</ref> 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.<ref>{{cite web|url=http://www.sbs.com.au/news/article/2016/05/11/drivers-pose-significant-threat-endangered-tasmanian-devil|title=Drivers pose 'significant' threat to endangered Tasmanian devil|work=News|date=14 May 2016 |access-date=14 May 2016}}</ref><ref>{{cite web|url=http://www.abc.net.au/news/2015-12-09/alarm-system-to-reduce-tasmanian-devil-roadkill-shows-promise/7013558|title='Virtual fence' shows promise in reducing road toll of Tasmanian devils|date=9 December 2015|work=ABC News|access-date=14 May 2016}}</ref> ===Devil facial tumour disease=== {{Main|Devil facial tumour disease}} [[File:Tasmanian Devil Facial Tumour Disease.png|thumb|Devil facial tumour disease causes tumours to form in and around the mouth, interfering with feeding and eventually leading to death by starvation.|alt=A black devil with several pink-red coloured tumours growing on many parts of its body. The two largest are one covering where its right eye is, and another below the left eye. The right eye is no longer visible and both of these are around one-third the size of a normal devil face. It is lying on a green fabric.]] 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.<ref name=Deakin2012/><ref name=DFTDUpdateJune20005>{{cite web|publisher=Department of Primary Industries, Parks, Water and Environment |date=June 2005 |url=http://www.dpiwe.tas.gov.au/inter.nsf/Attachments/LBUN-6D73V5/$FILE/Tas_devil_update_June2005.pdf |title=Devil Facial Tumour Disease Update |access-date=30 September 2010 |url-status=dead |archive-url=https://web.archive.org/web/20080907075720/http://www.dpiwe.tas.gov.au/inter.nsf/Attachments/LBUN-6D73V5/%24FILE/Tas_devil_update_June2005.pdf |archive-date=7 September 2008 }}</ref><ref name=DPIWEDMS2005>{{cite web|publisher=Department of Primary Industries, Parks, Water and Environment |date=February 2005 |url=http://www.dpiwe.tas.gov.au/inter.nsf/Attachments/LBUN-6996MH/$FILE/DFTD_DMS_Feb05a.pdf |title=Tasmanian Devil Facial Tumour Disease (DFTD) Disease Management Strategy |access-date=30 September 2010 |url-status=dead |archive-url=https://web.archive.org/web/20080907075712/http://www.dpiwe.tas.gov.au/inter.nsf/Attachments/LBUN-6996MH/%24FILE/DFTD_DMS_Feb05a.pdf |archive-date=7 September 2008 }}</ref> Individual devils die within months of infection.<ref name=DPIWEweb3>{{cite web|publisher=Department of Primary Industries, Parks, Water and Environment |url=http://www.dpiwe.tas.gov.au/inter.nsf/WebPages/LBUN-5QF86G?open |title=Devil Facial Tumour Disease |access-date=30 September 2010 |url-status=dead |archive-url=https://web.archive.org/web/20050921014142/http://www.dpiwe.tas.gov.au/inter.nsf/WebPages/LBUN-5QF86G?open |archive-date=21 September 2005 }}</ref> The disease is an example of [[transmissible cancer]], which means that it is contagious and passed from one animal to another.<ref name=shea>{{cite news|author=Shea, N.|title=Wildlife: Devils in danger|work=National Geographic|date=November 2006}}</ref> This tumour is able to pass between hosts without inducing a response from the host's immune system.<ref name="Siddle 16221–16226">{{Cite journal|last1=Siddle|first1=Hannah V.|last2=Kreiss|first2=Alexandre|last3=Eldridge|first3=Mark D. B.|last4=Noonan|first4=Erin|last5=Clarke|first5=Candice J.|last6=Pyecroft|first6=Stephen|last7=Woods|first7=Gregory M.|last8=Belov|first8=Katherine|date=2007-10-09|title=Transmission of a fatal clonal tumor by biting occurs due to depleted MHC diversity in a threatened carnivorous marsupial|journal=Proceedings of the National Academy of Sciences|language=en|volume=104|issue=41|pages=16221–16226|doi=10.1073/pnas.0704580104|issn=0027-8424|pmc=1999395|pmid=17911263|doi-access=free}}</ref> Dominant devils who engage in more biting behaviour are more exposed to the disease.<ref>{{cite journal |last1=Wells |first1=Konstans |last2=Hamede |first2=Rodrigo |last3=Kerlin |first3=Douglas |last4=Storfer |first4=Andrew |last5=Hohenlohe |first5=Paul |last6=Jones |first6=Menna |last7=McCallum |first7=Hamish |title=Infection of the fittest: devil facial tumour disease has greatest effect on individuals with highest reproductive output |journal=Ecology Letters |date=2017 |volume=20 |issue=6 |pages=770–778 |doi=10.1111/ele.12776 |pmid=28489304 |pmc=6759051 |bibcode=2017EcolL..20..770W |url=https://cronfa.swan.ac.uk/Record/cronfa45075 }}</ref> 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.<ref name=DPIWEDMS2005/> 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.<ref name=DPIWEDMS2005/> In March 2017, scientists at the University of Tasmania presented an apparent first report of having successfully treated Tasmanian devils with the disease. Live cancer cells that were treated with [[IFN-γ]] to restore [[MHC-I]] expression, were injected into the infected devils to stimulate their immune system to recognise and fight the disease.<ref>{{cite journal |vauthors=Tovar C, Pye RJ, Kreiss A, Cheng Y, Brown GK, Darby J, Malley RC, Siddle HV, Skjødt K, Kaufman J, Silva A, Baz Morelli A, Papenfuss AT, Corcoran LM, Murphy JM, Pearse MJ, Belov K, Lyons AB, Woods GM |display-authors=6 |title=Regression of devil facial tumour disease following immunotherapy in immunised Tasmanian devils |journal=Scientific Reports |volume=7 |pages=43827 |date=March 2017 |pmid=28276463 |pmc=5343465 |doi=10.1038/srep43827|bibcode=2017NatSR...743827T }}</ref> In 2020 it was reported that one of the last DFTD-free wild population of Tasmanian devils was suffering from inbreeding depression and has undergone a significant decline in reproductive success in recent years.<ref>{{cite journal | last1 = Gooley | first1 = RM | last2 = Hogg | first2 = CJ | last3 = Fox | first3 = S | last4 = Pemberton | first4 = D | last5 = Belov | first5 = K | last6 = Grueber | first6 = CE | year = 2020 | title = Inbreeding depression in one of the last DFTD-free wild populations of Tasmanian devils | journal = PeerJ | volume = 8 | issue = | page = e9220 | doi = 10.7717/peerj.9220 | doi-access = free | pmid = 32587794 | pmc = 7304431 }}</ref> ==Relationship with humans== At [[Lake Nitchie]] in western [[New South Wales]] in 1970, a male human skeleton wearing a necklace of 178 teeth from 49 different devils was found. The skeleton is estimated to be 7000 years old, and the necklace is believed to be much older than the skeleton. Archaeologist [[Josephine Flood]] believes the devil was hunted for its teeth and that this contributed to its extinction on mainland Australia. Owen and Pemberton note that few such necklaces have been found.<ref>Owen and Pemberton, p. 42.</ref> [[Midden]]s that contain devil bones are rare—two notable examples are [[Devil's Lair]] in the south-western part of Western Australia and [[Tower Hill State Game Reserve|Tower Hill]] in Victoria.<ref>Owen and Pemberton, p. 41.</ref> In Tasmania, [[Tasmanian Aboriginals|local Indigenous Australians]] and devils sheltered in the same caves. Tasmanian Aboriginal names for the devil recorded by Europeans include "tarrabah", "poirinnah", and "par-loo-mer-rer".<ref>Owen and Pemberton, p. 3.</ref> Variations also exist, such as "Taraba" and "purinina".<ref>{{Cite web|title=Taraba : Tasmanian Aboriginal Stories|url=https://www.ncacl.org.au/atsi_resource/taraba-tasmanian-aboriginal-stories/|access-date=2020-10-07|website=NCACL|language=en}}</ref><ref>{{Cite web|date=2017-08-15|title=Native animals should be rechristened with their Aboriginal names|url=https://www.australiangeographic.com.au/topics/wildlife/2017/08/native-animals-should-be-renamed-with-their-aboriginal-names/|access-date=2020-10-07|website=Australian Geographic|language=en-AU}}</ref> It is a common belief that devils will eat humans. While they are known to eat dead bodies, there are prevalent myths that they eat living humans who wander into the bush.<ref>Owen and Pemberton, p. 10.</ref> Despite outdated beliefs and exaggerations regarding their disposition, many, although not all, devils will remain still when in the presence of a human; some will also shake nervously. They can bite and scratch out of fear when held by a human, but a firm grip will cause them to remain still.<ref>Owen and Pemberton, pp. 15–18.</ref> Although they can be tamed, they are asocial, and are not considered appropriate as pets;<ref name="Owen and Pemberton-25"/> they have an unpleasant odour, and neither demonstrate nor respond to affection.<ref>Owen and Pemberton, p. 113.</ref> Until recently, the devil was not studied much by academics and naturalists.<ref name="Owen and Pemberton-4"/> At the start of the 20th century, Hobart zoo operator Mary Roberts, who was not a trained scientist, was credited for changing people's attitudes and encouraging scientific interest in native animals (such as the devil) that were seen as fearsome and abhorrent, and the human perception of the animal changed.<ref name=orob/> [[Theodore Thomson Flynn]] was the first professor of biology in Tasmania, and carried out some research during the period around World War I.<ref>Owen and Pemberton, pp. 93–97.</ref> In the mid-1960s, Professor Guiler assembled a team of researchers and started a decade of systematic fieldwork on the devil. This is seen as the start of modern scientific study of it.<ref>Owen and Pemberton, pp. 99–101.</ref> However, the devil was still negatively depicted, including in tourism material.<ref name="Owen and Pemberton-99"/> The first doctorate awarded for research into the devil came in 1991.<ref name="Owen and Pemberton-4">Owen and Pemberton, p. 4.</ref> ===In captivity=== [[File:Sarcophilus harrisii -Healesville Sanctuary-8a.jpg|thumb|At [[Healesville Sanctuary]], Victoria|alt=A devil with red ears and white patches under its neck, is standing on some bark chips, in front of some grass and behind a rock of the size of its body.]] 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]].<ref name=orob>Owen and Pemberton, pp. 84–93.</ref> Even by 1934, successful breeding of the devil was rare.<ref>Owen and Pemberton, pp. 67–69.</ref> 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.<ref name=Phillips2003/> In general, females tend to retain more stress after being taken into captivity than males.<ref name=Jones2005/> Tasmanian devils were displayed in various zoos around the world from the 1850s onwards.<ref>Owen and Pemberton, p. 132.</ref> In the 1950s several animals were given to European zoos.<ref>Owen and Pemberton, pp. 101–2.</ref> In October 2005 the Tasmanian government sent four devils, two male and two female, to the [[Copenhagen Zoo]], following the birth of the [[Christian, Crown Prince of Denmark|first son]] of [[Frederik X|King Frederik X of Denmark]] and his Tasmanian-born wife [[Queen Mary of Denmark|Mary]].<ref>{{cite news|url=http://www.smh.com.au/news/world/marys-little-devils/2006/04/11/1144521306868.html|date=11 April 2006|access-date=14 September 2010|work=The Sydney Morning Herald|title=Mary's little devils}}</ref> Due to restrictions on their export by the Australian government, at the time these were the only devils known to be living outside Australia.<ref name=fed/> 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.<ref>{{cite web|title=Ambassador Devils for Overseas Zoos|url=http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/25ED286F84111687CA257B940000CBF4?OpenDocument|publisher=Save the Tasmanian Devil|access-date=30 November 2014|archive-date=13 March 2018|archive-url=https://web.archive.org/web/20180313125019/http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/25ED286F84111687CA257B940000CBF4?OpenDocument|url-status=dead}}</ref> [[San Diego Zoo Wildlife Alliance]] and [[Albuquerque Biological Park|Albuquerque Biopark]] were selected to participate in the program,<ref>{{cite web|title=First overseas zoos selected for ambassador devils |url=http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/204F7A6644F6B28DCA257BD9007E6911 |publisher=Save the Tasmanian Devil |access-date=30 November 2014 |url-status=dead |archive-url=https://web.archive.org/web/20150904025041/http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/204F7A6644F6B28DCA257BD9007E6911 |archive-date=4 September 2015 }}</ref> and [[Wellington Zoo]] and [[Auckland Zoo]] soon followed.<ref>{{cite web|title=Auckland Zoo helps raise awareness of Tasmanian devils |url=http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/219544C610F00ACBCA257CD6000520C8 |publisher=Save the Tasmanian Devil |access-date=30 November 2014 |url-status=dead |archive-url=https://web.archive.org/web/20150904025041/http://www.tassiedevil.com.au/tasdevil.nsf/TheProgram/219544C610F00ACBCA257CD6000520C8 |archive-date=4 September 2015 }}</ref> 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]],<ref>{{Cite web|url=http://kidszoo.org/tasmanian-devils-returning-to-zoo/|title=Tasmanian Devils Returning to Zoo|date=January 27, 2015|website=kidszoo.org|publisher=Fort Wayne Children's Zoo|access-date=2016-07-24|archive-url=https://web.archive.org/web/20160603224256/http://kidszoo.org/tasmanian-devils-returning-to-zoo/|archive-date=3 June 2016|url-status=dead|df=dmy-all}}</ref> the [[Los Angeles Zoo]],<ref>{{Cite web|url=http://www.lazoo.org/2015/12/tasmanian-devils/|title=Tasmanian Devils are Back at the L.A. Zoo After 20 Years!|date=December 14, 2015|website=lazoo.org|publisher=Los Angeles Zoo and Botanical Gardens|access-date=July 24, 2016|archive-date=20 March 2020|archive-url=https://web.archive.org/web/20200320132142/https://www.lazoo.org/2015/12/tasmanian-devils/|url-status=dead}}</ref> the [[Saint Louis Zoo]],<ref>{{Cite web|url=http://www.stltoday.com/news/local/metro/new-to-the-st-louis-zoo-tasmanian-devils/article_eb49ad3d-1382-5228-ad8f-4ca0f8ae2e19.html|title=New to the St. Louis Zoo: Tasmanian devils|last=Bock|first=Jessica|date=April 20, 2016|website=stltoday.com|publisher=St. Louis Post-Dispatch|access-date=2016-07-24}}</ref> and the [[Toledo Zoo]].<ref>{{Cite web|url=http://www.toledoblade.com/local/2015/09/06/Toledo-Zoo-joins-effortto-save-Tasmanian-devils.html|title=Toledo Zoo joins effort to save Tasmanian devils|last=Mester|first=Alexandra|date=September 6, 2015|website=The Blade|access-date=2016-07-24}}</ref> Captive devils are usually forced to stay awake during the day to cater to visitors, rather than following their natural nocturnal style.<ref>Owen and Pemberton, p. 133.</ref> ===In popular culture=== [[File:Taz Looney Tunes parade.jpg|right|thumb|Warner Bros' Tasmanian Devil "Taz" at a parade in California|alt=A black open-top sports car with lights on is being driven down an asphalt road. A large furry toy costume, slightly larger than a human is standing in the back seat. It has cream coloured mouth and chest, and dark brown arms and forehead, large whiskers, a grin, large white eyes and two canines. Behind him are some men walking in green costumes. On the left is a crowd watching the parade from the footpath, in front of tall buildings with stone arches.]] 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]],<ref name=DPIWEweb1/> and the former Tasmanian Australian rules football team which played in the [[Victorian Football League]] was known as the [[Tasmania Devils Football Club (VFL)|Devils]].<ref>{{Cite news|url=http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/DevilNews_June_2008.pdf|date=June 2008|access-date=6 October 2010|work=Save the Tasmanian Devil|title=Welcome|page=1|archive-date=17 February 2011|archive-url=https://web.archive.org/web/20110217185140/http://www.tassiedevil.com.au/tasdevil.nsf/downloads/D595436FECB69A66CA2576ED0083D3F6/$file/DevilNews_June_2008.pdf|url-status=dead}}</ref> The [[Hobart Devils]] were once part of the [[National Basketball League (Australia)|National Basketball League]].<ref>{{cite web |url=http://www.nbl.com.au/award-winners/ |title=Most Valuable Player |publisher=[[National Basketball League (Australasia)|National Basketball League]] |access-date=4 September 2015 |url-status=dead |archive-url=https://web.archive.org/web/20151222082507/http://www.nbl.com.au/award-winners/ |archive-date=22 December 2015 |df=dmy-all }}</ref> The devil has appeared on several [[Coins of the Australian dollar#Collectable coins|commemorative coins]] in Australia over the years.<ref>{{cite web|url=http://www.perthmint.com.au/catalogue/2009-celebrate-australia-1-coin-tasmania.aspx|publisher=The Perth Mint|title=2009 Celebrate Australia $1 coin – Tasmania|access-date=6 October 2010|archive-date=10 October 2010|archive-url=https://web.archive.org/web/20101010000538/http://www.perthmint.com.au/catalogue/2009-celebrate-australia-1-coin-tasmania.aspx|url-status=dead}}</ref><ref>{{cite web|url=http://mintissue.ramint.gov.au/mintissue/product.asp?code=802152 |publisher=Royal Australian Mint |title=2010 $5 Gold Proof Tinga Tasmanian Devil |access-date=6 October 2010 |url-status=dead |archive-url=https://web.archive.org/web/20100813182845/http://mintissue.ramint.gov.au/mintissue/product.asp?code=802152 |archive-date=13 August 2010 }}</ref> [[Cascade Brewery]] in Tasmania sells a [[ginger beer]] with a Tasmanian devil on the label.<ref>{{cite web|url=http://www.fostersgroup.com/brands/cascade-ginger-beer.aspx |title=Cascade Ginger Beer |access-date=6 October 2010 |publisher=[[Foster's Group]] |url-status=dead |archive-url=https://web.archive.org/web/20100609022137/http://www.fostersgroup.com/brands/cascade-ginger-beer.aspx |archive-date=9 June 2010 }}</ref> In 2015, the Tasmanian devil was chosen as Tasmania's [[Symbols of Tasmania|state emblem]].<ref>{{cite news|title=Tasmania backs the devil as the state emblem despite endangered status|url=http://www.abc.net.au/news/2015-05-31/tasmania-backs-the-devil-as-the-state-emblem/6510228|access-date=31 May 2015|work=ABC News|date=31 May 2015}}</ref> 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".<ref>{{cite web|url=http://www.tasmaniandevilpark.com/news.html |title=World tourism can help save the Tasmanian Devil, park director tells international conference |date=5 June 2008 |access-date=6 October 2010 |publisher=Tasmanian Devil Conservation Park |url-status=dead |archive-url=https://web.archive.org/web/20090914043540/http://www.tasmaniandevilpark.com/news.html |archive-date=14 September 2009 }}</ref> There has also been a multimillion-dollar proposal to build a giant 19 m-high, 35 m-long devil in [[Launceston, Tasmania|Launceston]] in northern Tasmania as a tourist attraction.<ref>{{cite news|publisher=Australian Broadcasting Corporation|url=http://www.abc.net.au/worldtoday/content/2006/s1561675.htm|author=Jeanes, Tim|title=Giant Tassie Devil tourist attraction in danger|date=3 February 2006|access-date=6 October 2010}}</ref> 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.<ref>Owen and Pemberton, pp. 122–124.</ref> The Tasmanian devil is probably best known internationally as the inspiration for the ''Looney Tunes'' cartoon character [[Tasmanian Devil (Looney Tunes)|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.<ref>Owen and Pemberton, p. 12.</ref> 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.<ref>Owen and Pemberton, pp. 156–160.</ref> 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.<ref>Owen and Pemberton, pp. 161–164.</ref> [[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.<ref>Owen and Pemberton, pp. 167, 169.</ref> In 2006, Warner Bros. permitted the Government of Tasmania to sell stuffed toys of Taz with profits funnelled into research on DFTD.<ref>{{Cite news|url=http://www.smh.com.au/news/National/Warner-Bros-to-help-save-Tassie-devils/2006/06/20/1150701535616.html|title=Warner Bros to help save Tassie devils|work=The Sydney Morning Herald|date=20 June 2006|access-date=30 September 2010}}</ref> ==See also== {{Portal|Mammals}} *[[Fauna of Australia]] *[[Threatened fauna of Australia]] *[[List of adaptive radiated marsupials by form]] ==References== <!-- BiolConserv131:307. --> ===Notes=== {{Reflist|refs= <ref name=Harris1807>{{cite journal|last=Harris|first=G. 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E. |author2=Paetkau, David |author3=Geffen, Eli |author4= Moritz, Craig |title=Genetic diversity and population structure of Tasmanian devils, the largest marsupial carnivore |journal=Molecular Ecology |year=2004 |volume=13 |issue=8 |pmid=15245394 |pages=2197–2209 |doi=10.1111/j.1365-294X.2004.02239.x|bibcode=2004MolEc..13.2197J |s2cid=14068282 }}</ref> <ref name="morris&austin2012">{{cite journal |last1=Morris |first1=K. |last2=Austin |first2=J. J. |last3=Belov |first3=K. |title=Low major histocompatibility complex diversity in the Tasmanian devil predates European settlement and may explain susceptibility to disease epidemics |journal=[[Biology Letters]] |date=5 December 2012 |volume=9 |issue=1 |pages=20120900 |doi=10.1098/rsbl.2012.0900|pmid=23221872 |pmc=3565505 }}</ref> <ref name=LachishMiller2010>{{Cite journal |last1=Lachish |first1= S. |last2=Miller |first2=K. |last3=Storfer |first3=A. |last4=Goldizen |first4=A. |last5=Jones |first5=M. |title=Evidence that disease-induced population decline changes genetic structure and alters dispersal patterns in the Tasmanian devil |journal=Heredity |volume=106 |issue=1 |pages=172–182 |year=2010 |pmid=20216571 |pmc=3183847 |doi=10.1038/hdy.2010.17 |display-authors=4}}</ref> <ref name=sid>{{cite journal |journal=Proceedings of the Royal Society B |year=2010 |volume=277 |author1=Siddle, Hannah V. |author2=Marzec, Jolanta |author3=Cheng, Yuanyuan |author4=Jones, Menna |author5=Belov, Katherine |title=MHC gene copy number variation in Tasmanian devils: Implications for the spread of a contagious cancer |doi=10.1098/rspb.2009.2362 |pages=2001–06 |pmid=20219742 |pmc=2880097 |issue=1690 |display-authors=4}}</ref> <ref name=Jones2008>{{cite journal|last=Jones |first=M. E. |author2=Cockburn, A. |author3=Hamede, R |author4=Hawkins, C. |author5=Hesterman, H. |author6=Lachish, S. |author7=Mann, D. |author8=McCallum, H. |author9=Pemberton, D. |title=Life-history change in disease-ravaged Tasmanian devil populations |journal=[[Proceedings of the National Academy of Sciences]] |volume=105 |issue=29 |pages=10023–7 |year=2008 |doi=10.1073/pnas.0711236105 |pmc=2481324 |pmid=18626026 |display-authors=4|bibcode=2008PNAS..10510023J |doi-access=free }}</ref> <ref name=Wroe2005>{{cite journal |author1=Wroe, S. |author2=McHenry, C. |author3=Thomason, J. |year=2005 |title=Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa |journal=Proceedings of the Royal Society B: Biological Sciences |volume=272 |pages=619–625 |pmid=15817436 |pmc=1564077 |doi=10.1098/rspb.2004.2986 |issue=1563}}</ref> <ref name=Wroe1999>{{cite journal|doi=10.1111/1475-4983.00082 |author=Wroe, Stephen |title=The geologically oldest dasyurid, from the Miocene of Riversleigh, north-west Queensland |journal=Palaeontology |volume=42 |issue=3 |year=1999 |pages=501–527|doi-access=free |bibcode=1999Palgy..42..501W }}</ref> <ref name=jb98>{{cite journal|doi=10.1046/j.1365-2656.1998.00203.x|author1=Jones, Menna E. |author2=Barmuta, Leon A. |year=1988 |title=Diet overlap and relative abundance of sympatric dasyurid carnivores: a hypothesis of competition |journal=Journal of Animal Ecology |volume=67 |pages=410–421 |issue=3|doi-access=free }}</ref> <ref name=Hamede>{{cite journal|title=Contact networks in a wild Tasmanian devil (''Sarcophilus harrisii'') population: using social network analysis to reveal seasonal variability in social behaviour and its implications for transmission of devil facial tumour disease|author1=Hamede, Rodrigo K. |author2=Bashford, Jim |author3=McCallum, Hamish |author4=Jones, Menna E. |journal=Ecology Letters |volume=12 |issue=11 |pages=1147–57 |date=November 2009 |pmid=19694783 |doi=10.1111/j.1461-0248.2009.01370.x|bibcode=2009EcolL..12.1147H |url=https://figshare.com/articles/journal_contribution/22871279 }}</ref> <ref name=Cooper2010>{{cite journal|title=Comparative physiology of Australian quolls (''Dasyurus''; Marsupialia)|author1=Cooper, Christine E. |author2=Withers, Philip C.|journal=Journal of Comparative Physiology B |volume=180 |issue=6 |pages=857–68 |year=2010 |doi=10.1007/s00360-010-0452-3 |pmid=20217094|url=https://espace.curtin.edu.au/bitstream/20.500.11937/8095/2/147157_147157.pdf|hdl=20.500.11937/8095 |s2cid=7440785 |hdl-access=free }}</ref> <ref name=Pemberton1993>{{Cite journal|doi=10.1071/ZO9930507|author=Pemberton, David |author2=Renouf, Deane |year=1993 |title=A field-study of communication and social behaviour of Tasmanian Devils at feeding sites |journal=Australian Journal of Zoology |volume=41 |pages=507–26 |issue=5}}</ref> <ref name=jb00>{{cite journal|doi=10.1644/1545-1542(2000)081<0434:NDASAD>2.0.CO;2 |author=Jones, Menna E. |author2=Barmuta, Leon A. |year=2000 |title=Niche differentiation among sympatric Australian dasyurid carnivores |journal=Journal of Mammalogy |volume=81 |pages=434–47 |issue=2|doi-access=free }}</ref> <ref name=Guiler1970>{{cite journal |doi=10.1071/ZO9700063 |author=Guiler, E. 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N. |author2=Wroe, S. |year=2003 |title=Causes of extinction of vertebrates during the Holocene of mainland Australia: arrival of the dingo, or human impact? |journal=Holocene |volume=13 |pages=941–948 |doi=10.1191/0959683603hl682fa |issue=6|bibcode=2003Holoc..13..941J |s2cid=15386196 }}</ref> <ref name=Bradshaw2005>{{cite journal|doi=10.1111/j.0906-7590.2005.04088.x |author=Bradshaw, C. J. A. |author2=Brook, B. W. |year=2005 |title=Disease and the devil: density-dependent epidemiological processes explain historical population fluctuations in the Tasmanian devil |journal=Ecography |volume=28 |pages=181–190 |issue=2|bibcode=2005Ecogr..28..181B }}</ref> <ref name="distimpactdftd">{{Cite journal |last1=McCallum |first1=H. |last2=Tompkins |first2=D. M. |last3=Jones |first3=M. |last4=Lachish |first4=S. |last5=Marvanek |first5=S. |last6=Lazenby |first6=B. |last7=Hocking |first7=G. |last8=Wiersma |first8=J. |last9=Hawkins |first9=C. E. |title=Distribution and Impacts of Tasmanian Devil Facial Tumor Disease |journal=EcoHealth |volume=4 |issue=3 |pages=318–325 |year=2007 |doi=10.1007/s10393-007-0118-0 |citeseerx=10.1.1.464.5369 |s2cid=11311742 |display-authors=4 |url=http://fpop.fpa.tas.gov.au/staffPublications/2007_devil_disease.pdf |access-date=27 October 2017 |archive-date=1 March 2015 |archive-url=https://web.archive.org/web/20150301124347/http://fpop.fpa.tas.gov.au/staffPublications/2007_devil_disease.pdf |url-status=dead }}</ref> <ref name=LachishMcCallum>{{cite journal |last1=Lachish |first1=Shelly |last2=McCallum |first2=Hamish |last3=Mann |first3=Dydee |last4=Pukk |first4=Chrissy E. |last5=Jones |first5=Menna E. |title=Evaluation of Selective Culling of Infected Individuals to Control Tasmanian Devil Facial Tumor Disease |journal=[[Conservation Biology (journal)|Conservation Biology]] |date=19 January 2010 |volume=24 |issue=3 |pages=841–851 |doi=10.1111/j.1523-1739.2009.01429.x |display-authors=4 |pmid=20088958|s2cid=13424807 |doi-access=free |bibcode=2010ConBi..24..841L }}</ref> <ref name=BeetonMcCallum>{{cite journal|last1=Beeton|first1=Nick|last2=McCallum|first2=Hamish|title=Models predict that culling is not a feasible strategy to prevent extinction of Tasmanian devils from facial tumour disease|journal=[[Journal of Applied Ecology]]|date=December 2011|volume=48|issue=6|pages=1315–1323|doi=10.1111/j.1365-2664.2011.02060.x|doi-access=free|bibcode=2011JApEc..48.1315B }}</ref> <ref name="Hobday-2010">{{Cite journal |last1=Hobday |first1=Alistair J. |title=Nighttime driver detection distances for Tasmanian fauna: informing speed limits to reduce roadkill |journal=Wildlife Research |volume=37 |pages=265–272 |year=2010 |doi=10.1071/WR09180 |issue=4|bibcode=2010WildR..37..265H }}</ref> <ref name="Jones, Menna E.-2000">{{Cite journal |doi=10.1071/WR98069 |title=Road upgrade, road mortality and remedial measures: impacts on a population of eastern quolls and Tasmanian devils |author=Jones, Menna E. |journal=Wildlife Research |volume=27 |year=2000 |pages=289–296 |issue=3|bibcode=2000WildR..27..289J }}</ref> <ref name=Deakin2012>{{cite journal |journal=Annual Review of Genomics and Human Genetics |title=A Comparative Genomics Approach to Understanding Transmissible Cancer in Tasmanian Devils |volume=13 |year=2012 |pages=207–222 |doi=10.1146/annurev-genom-090711-163852 |last1=Deakin |first1=Janine E. |last2=Belov |first2=Katherine |pmid=22657390}}</ref> <ref name=Phillips2003>{{Cite journal|doi=10.1002/zoo.10092 |author=Phillips, B. T. |author2=Jackson, S. M. |year=2003 |title=Growth and development of the Tasmanian devil (Sarcophilus harrisii) at Healesville Sanctuary, Victoria, Australia |journal=Zoo Biology |volume=22 |pages=497–505 |issue=5}}</ref> <ref name=Jones2005>{{cite journal |doi=10.1071/ZO05043 |author=Jones, S. M. |author2=Lockhart, T. J. |author3=Rose, R. W. |year=2005 |url=http://www.zoo.utas.edu.au/RR/pdfRose/Devil.pdf |title=Adaptation of wild-caught Tasmanian devils (Sarcophilus harrisii) to captivity: evidence from physical parameters and plasma cortisol concentrations |journal=Australian Journal of Zoology |volume=53 |pages=339–344 |issue=5 |access-date=1 September 2010 |archive-date=4 March 2011 |archive-url=https://web.archive.org/web/20110304205635/http://www.zoo.utas.edu.au/RR/pdfRose/Devil.pdf |url-status=dead }}</ref> }} ===Bibliography=== {{Refbegin}} *{{Cite book|last=Guiler|first=ER|title=The Tasmanian devil|year=1992|publisher=St David's Park Publishing|isbn=0-7246-2257-8|location=Hobart, Tasmania}} *{{cite book|title=The Transformers Beast Wars Sourcebook|year=2008|location=San Diego, California|publisher=[[IDW Publishing]]|isbn=978-1-60010-159-5 |author=Figueroa, Don |author2=Furman, Simon |author3=Yee, Ben |author4=Khanna, Dan M. |author5=Guidi, Guido |author6=Isenberg, Jake |author7=Matere, Marcelo |author8=Roche, Roche |author9=Ruffolo, Rob |author10=Williams, Simon}} *{{Cite book|last=Owen|first=D|title=Tasmanian Devil: A unique and threatened animal|year=2005|publisher=[[Allen & Unwin]]|isbn=978-1-74114-368-3|url=https://books.google.com/books?id=VrhlpKtUnu8C&q=Tasmanian+Devil|author2=Pemberton, David |access-date=22 August 2010|location=Crows Nest, New South Wales}} *{{cite book|author=Paddle, Robert|year=2000|title=The Last Tasmanian Tiger: The History and Extinction of the Thylacine|location=Oakleigh, Victoria|publisher=Cambridge University Press|isbn=0-521-53154-3}} *{{Cite book|author=Tyndale-Biscoe, Hugh|title=Life of marsupials|location=Collingwood, Victoria|publisher=CSIRO Publishing|year=2005|isbn=0-643-06257-2}} {{Refend}} ==Further reading== *{{cite journal |doi=10.1111/j.1469-7998.2008.00419.x |author=Hesterman, H. |author2=Jones, S. M. |author3=Schwarzenberger, F. |year=2008 |title=Pouch appearance is a reliable indicator of the reproductive status in the Tasmanian devil and the spotted-tailed quoll |journal=Journal of Zoology |volume=275 |pages=130–138 |issue=2|url=https://figshare.com/articles/journal_contribution/22867187 }} *{{Cite journal |last1=McDonald-Madden |first1=E. |last2=Probert |first2=W. J. M. |last3=Hauser |first3=C. E. |last4=Runge |first4=M. C. |last5=Possingham |first5 = H. P. |last6=Jones |first6=M. E. |last7=Moore |first7=J. L. |last8=Rout |first8=T. M. |last9=Vesk |first9=P. A. |last10=Wintle |first10=B. A. |title=Active adaptive conservation of threatened species in the face of uncertainty |journal=Ecological Applications |volume=20 |issue=5 |pages=1476–1489 |year=2010 |pmid=20666263 |doi=10.1890/09-0647.1|bibcode=2010EcoAp..20.1476M |url=http://espace.library.uq.edu.au/eserv/UQ:207388/UQ207388OA.pdf }} ==External links== {{Commons and category|Sarcophilus harrisii|Sarcophilus_harrisii}} {{Wikispecies|Sarcophilus harrisii}} {{Spoken Wikipedia|Tasmanian_Devil.ogg|date=2006-08-08}} *[https://web.archive.org/web/20080721030213/http://www.parks.tas.gov.au/wildlife/mammals/devil.html Parks and Wildlife Tasmania – Tasmanian Devil] – vocalisation, movie, FAQ (archived 21 July 2008) *[http://www.tassiedevil.com.au/ Save the Tasmanian Devil] {{Webarchive|url=https://web.archive.org/web/20090321072257/http://www.tassiedevil.com.au/ |date=21 March 2009 }} – Tasmanian government conservation program *View the [http://www.ensembl.org/Sarcophilus_harrisii Tasmanian devil genome] in [[Ensembl]] * {{UCSC genomes|sarHar1}} * [https://www.aussieark.org.au/devil-ark/ The Aussie Devil Ark Conservation Project] {{Dasyuromorphia|D.D.}} {{Taxonbar|from=Q134964}} {{Authority control}} [[Category:Mammals described in 1841]] [[Category:Dasyuromorphs]] [[Category:Mammals of Tasmania]] [[Category:Mammals of New South Wales]] [[Category:Marsupials of Australia]] [[Category:Scavengers]] [[Category:Vulnerable fauna of Australia]] [[Category:Species that are or were threatened by disease]] [[Category:Species that are or were threatened by collisions with vehicles]] [[Category:Symbols of Tasmania]]
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