Dingo

Domestic status

The dingo is regarded as a feral dog because it descended from domesticated ancestors.^[1][7] The dingo's relationship with indigenous Australians can be described as commensalism – "the process by which two organisms live in close association but they do not depend on each other for survival". They will both hunt and sleep together. The dingo is therefore comfortable enough around humans to associate with them, but is still capable of living independently.^[41] Any free-ranging unowned dog can be socialized to become an owned dog, as some dingoes do when they join human families.^[42] Although the dingo exists in the wild,^[14] it associates with humans but has not been selectively bred similarly to other domesticated animals.^[2][14] Therefore, its status as being a domestic animal is not clear.^[2] Whether the dingo was a wild or domesticated species was not clarified from Meyer's original description, which translated from the German language ambiguously reads:

It is not known if it is the only dog species in New South Wales, and if it can also still be found in the wild state; however, so far it appears to have lost little of its wild condition; moreover, no divergent varieties have been discovered.^[5]

Archaeological evidence

The oldest reliable date for dog remains found in mainland Southeast Asia is from Vietnam at 4,000 years before present (YBP), and in island southeast Asia from Timor-Leste at 3,000 YBP.^[43] The earliest dingo remains in the Torres Straits date to 2,100 YBP. In New Guinea, the earliest dog remains date to 2,500–2,300 YBP from Caution Bay near Port Moresby but no ancient New Guinea singing dog remains have been found.^[1]

The earliest dingo skeletal remains in Australia are estimated at 3,450 YBP from the Mandura Caves on the Nullarbor Plain, south-eastern Western Australia;^[1][2] 3,320 YBP from Woombah Midden near Woombah, New South Wales; and 3,170 YBP from Fromme's Landing on the Murray River near Mannum, South Australia.^[2] Dingo bone fragments were found in a rock shelter located at Mount Burr, South Australia in a layer that was originally dated 7,000-8,500 YBP.^[44] Excavations later indicated that the levels had been disturbed, and the dingo remains "probably moved to an earlier level."^[12][45] The dating of these early Australian dingo fossils led to the widely-held belief that dingoes first arrived in Australia 4,000 YBP and then took 500 years to disperse around the continent.^[14] However, the timing of these skeletal remains were based on the dating of the sediments in which they were discovered, and not the specimens themselves.^[43]

In 2018, the oldest skeletal bones from the Madura Caves were directly carbon dated between 3,348–3,081 YBP, providing firm evidence of the earliest dingo and that dingoes arrived later than had previously been proposed. The next most reliable timing is based on desiccated flesh dated 2,200 YBP from Thylacine Hole, 110 km west of Eucla on the Nullarbor Plain, south-eastern Western Australia. When dingoes first arrived they would have been taken up by Indigenous Australians, who then provided a network for their swift transfer around the continent. Based on the recorded distribution time for dogs across Tasmania and cats across Australia once Indigenous Australians had acquired them, the dispersal of dingoes from their point of landing until they occupied continental Australia is proposed to have taken only 70 years.^[43] The red fox is estimated to have dispersed across the continent in only 60–80 years.^[14]

Based on a comparison with these early fossils, dingo morphology has not changed over thousands of years. This suggests that there has been no artificial selection over this period and that the dingo represents an early form of dog.^[14] They have lived, bred, and undergone natural selection in the wild, isolated from other canines until the arrival of European settlers, resulting in a unique canid.^[22][16]

Lineage

The Sahul Shelf and the Sunda Shelf during the past 12,000 years. Tasmania separated from the mainland 12,000 YBP,^[46] New Guinea separated from the mainland 6,500^[21]–8,500 YBP.^[21][47]

Proposed route for the migration of dogs based on mDNA. Haplotype A29 relates most to the Australian Dingo and the New Guinea Singing Dog, the ancient Polynesian Dog haplotype Arc2 to modern Polynesian, Indonesian and ancient New Zealand dogs, and the ancient Polynesian Arc1 is indistinguishable from a number of widespread modern haplotypes.^[1][48]

Whole genome sequencing has been used to survey "...the supposedly ancient, semi-domestic dog breeds, the Basenji and dingo."^[49] In 2016, a study based on whole genome sequences indicated that the dog is a genetically divergent subspecies of the gray wolf and was derived from a now-extinct ghost population of Late Pleistocene wolves, the dog and the dingo were not separate species,^[18] and in accordance with previous studies that the dingo and the Basenji are considered to be basal members of the domestic dog clade.^[18][19][20] "The term basal refers to a lineage that diverges early in the history of the group...and lies on a branch that originates near the common ancestor of the group."^[50] The study found evidence for gene flow of 0.3% mitochondrial DNA into the dingo lineage from the Qinghai population of the Tibetan wolf.^[18] The Australian National Kennel Council recognizes a dingo breed standard within its Hounds group.^[51]

At the end of the Last glacial maximum and the rise in sea levels, Tasmania became separated from the Australian mainland 12,000 YBP,^[46] and New Guinea 6,500^[21]–8,500 YBP^[21][47] by the inundation of the Sahul Shelf.^[52] No remains have been uncovered in Tasmania, therefore the Dingo is estimated to have arrived in Australia between 3,500-12,000 YBP. To reach Australia through the Malay Archipelago even at the lowest sea level of the Last Glacial Maximum, a journey of at least 50 km (30 mi) over open sea between ancient Sunda and Sahul was necessary, indicating that the dingo arrived to Sahul by boat.^[53]

A haplotype (haploid genotype) is a group of genes in an organism that are inherited together from a single parent.^[54][55] All dingo sequences studied exhibit mDNA haplotype A29, which falls within the Clade A haplogroup that represents 70% of domestic dogs.^{[56][57][58][53][59]} The evidence suggests that the haplotype was introduced from East Asia^[53] or southeast Asia^[56][60] through the islands of the Malay archipelago and into Australia.^[53][56][60] Haplotype A29 was one of several domestic dog mDNA haplotypes brought into the Malay Archipelago but only A29 reached mainland Australia.^[53] The mDNA haplotype A29, or a haplotype one mutation step away, was found in all of the Australian dingoes and New Guinea Singing Dogs so far studied, indicating descent from a common female ancestor.^[56]

In 2011, a study of the dingo male lineage using Y chromosome DNA (yDNA) as a genetic marker was undertaken for 338 Australian dingoes, New Guinea Singing Dogs, and village dogs from the Malay Archipelago. The Bali dogs support the arrival of their ancestors with the Austronesian expansion and the arrival of other domesticates 3-4,500 YBP. The data confirms that dingoes carry the unique yDNA haplogroup (H60) and it has been derived from yDNA haplogroup H5. Haplogroup H5 was not found in the village dogs from the Malay Archipelago but it is common in Taiwan. One H5 specimen from Taiwan clustered with one H60 from Australia with the indication of a common male ancestor 4-5,000 YBP and coincides with the expansion of the Daic people of Southern China. The conclusion is that there were 2 expansions of two types of dogs. Southern China produced the first ancient regional breeds 8,000 years ago. These were then dominated and replaced by a later explosive expansion of genetically diverse dogs that had been bred in South East Asia. If so, the dingo and the New Guinea Singing Dog, that pre-date the dogs of the Malay Archipelago, would reflect the last vestiges of the earlier ancient breeds.^[61]

The existence of a genetic subdivision within the dingo population has been proposed over two decades but has not been investigated.^[53][62] In 2016, a study compared DNA sequences using the entire mDNA genome (16,000 base pairs in length), and 13 DNA loci of the cell nucleus, taken from dingoes and New Guinea singing dogs. The study of their maternal mDNA provided evidence that they form a monophyletic clade (indicating that they all carried the same mutation inherited from a single female ancestor in the past). Dogs from China, Bali and Kalimantan did not fall within this clade. There are two distinct populations of dingoes in Australia based on both mitochondrial and nuclear evidence. The dingoes found today in the northwestern part of the Australian continent are estimated to have diverged 8,300 YBP, followed by a divergence of the New Guinea singing dog from the southeastern dingoes 7,800 YBP. As the New Guinea singing dog is closely related to the southeastern dingoes, these divergences are thought to have occurred somewhere in Sahul (a landmass which once included Australia, New Guinea and some surrounding islands)^[21] that existed until 6,500-8,000 years ago.^[21][14] The New Guinea singing dog then became a distinct but closely related lineage.^{[23][21][61][60]} The Fraser Island dingoes were unique because they cluster with the southeastern dingoes but exhibit many alleles (gene expressions) similar to the New Guinea singing dog, in addition to showing signs of admixture with the northwestern dingoes.^[21] These dates suggest that dingoes spread from Papua New Guinea to Australia over the land bridge at least twice.^[23][21] The lack of fossil evidence from northern Australia and Papua New Guinea could be explained by their tropical climate and acidic soil, as there are generally few fossils found in these regions.^[21] In 2017, a study of dingoes across a wider area found that the New Guinea singing dog female lineage is more closely related to the southeastern dingoes, and its male lineage is more closely related to dingoes found across the rest of the continent, indicating that the dingo lineage has a complex history.^[23]

The dingo and New Guinea singing dog lineage can be traced back through the Malay Archipelago to Asia. Therefore, the dingo is not a "wild dog" but a domesticated dog that has gone feral. Human-dog relationships are fluid, and a classification of being either completely "wild" or completely "domesticated" becomes difficult. They have adapted to surviving without human assistance and are the only known dog population to have maintained a long-term independence from humans.^[1]

Hybrids, distribution and habitat

Distribution map of dingoes and dingo-dog hybrids. The black line represents the Dingo Fence^[4]

The wolf-like canids are a group of large carnivores that are genetically closely related because their chromosomes number 78, therefore they can potentially interbreed to produce fertile hybrids.^[40] In the Australian wild there exists dingoes, feral dogs, and the crossings of these two which produces dingo-dog hybrids.^[4] Most studies looking at the distribution of dingoes focus on the distribution of dingo-dog hybrids instead.^[24]

Dingoes occurred throughout mainland Australia before European settlement.^[67][4] Dingoes are not found in the fossil record of Tasmania, therefore they arrived in Australia after Tasmania had separated from the mainland due to rising sea levels.^[68] The introduction of agriculture reduced dingo distribution, and by the early 1900s large barrier fences, including the Dingo Fence, excluded them from the sheep grazing areas. Land clearance, poisoning, and trapping caused the extinction of the dingo and hybrids from most of their former range in southern Queensland, New South Wales, Victoria and South Australia. Today, they are absent from most of New South Wales, Victoria, the south-eastern third of South Australia, and the south-western tip of Western Australia. They are sparse in eastern half of Western Australia and the adjoining areas of the Northern Territory and South Australia. They are regarded as common across the remainder of the continent.^[67][4]

The dingo could be considered an ecotype or an ecospecies which has adapted to Australia's unique environment.^[17] The dingo's present distribution covers a variety of habitats, including the temperate regions of eastern Australia, the alpine moorlands of the eastern highlands,the arid hot deserts of Central Australia, and the tropical forests and wetlands of Northern Australia.^[24] The occupation of, and adaption to, these habitats may have been assisted by their relationship with indigenous Australians.^[15]

Prey

Dingo with a fish on Fraser Island

A twenty-year study of the dingo's diet was conducted across Australia by the federal and state governments. These examined a total of 13,000 stomach contents and fecal samples.^[25] For the fecal samples, it was possible to determine the matching tracks of foxes and feral cats and not include these samples in the study, but it was impossible to distinguish between the tracks left by dingoes from those of dingo hybrids or feral dogs.^[69] The study found that these canines prey on 177 species represented by 72.3% mammals (71 species), 18.8% birds (53 species), 3.3% vegetation (seeds), 1.8% reptiles (23 species), and 3.8% insects, fish, crabs, and frogs (28 species).^[25][26][24] The relative proportions of prey is much the same across Australia, apart from more birds being eaten in the north and south-east coastal regions, and more lizards in Central Australia.^[25] Some 80% of the diet consisted of 10 species: red kangaroo, swamp wallaby, cattle, dusky rat, magpie goose, common brushtail possum, long-haired rat, agile wallaby, European rabbit, and the common wombat.^[70][26] Of the mammals eaten, 20% could be regarded as large.^[25]

However, the relative proportions of the size of prey mammals varied across regions. In the tropical coast region of the Northern Territory, agile wallabies, dusky rats and magpie geese formed 80% of the diet. In Central Australia, the rabbit has become a substitute for native mammals, and during droughts cattle carcasses provide most of the diet. On the Barkly Tableland, there are no rabbits nor does any native species dominate the diet, except for long-haired rats that form plagues every 9 years. In the Fortescue River region, the large red kangaroo and euro dominate the diet as there are few smaller mammals in this area. On the Nullarbor Plain, rabbits and red kangaroos dominate the diet, and twice as much rabbit eaten as red kangaroo. In the temperate mountains of eastern Australia, swamp wallaby and red-necked wallaby dominate the diet on the lower slopes and wombat on the higher slopes. Possums are commonly eaten here when found on the ground.^[25] In coastal regions dingoes patrol the beaches for washed-up fish, seals, penguins and other birds.^[26]

Dingoes drink approximately one litre of water each day in the summer and half a litre in winter. In arid regions during the winter, dingoes may live from the liquid in the bodies of their prey, as long as the number of prey is sufficient. In arid Central Australia, weaned pups draw most of their water from their food. There, regurgitation of water by the females for the pups was observed. During lactation, captive females have no higher need of water than usual, since they consume the urine and feces of the pups and therefore recycle the water and keep the den clean.^[26] Tracked dingoes in the Strzelecki Desert regularly visited water-points every 3–5 days, with two dingoes surviving 22 days without water during both winter and summer.^[71]

Hunting behaviour

Dingoes, dingo hybrids and feral dogs usually attack from the rear as they pursue their prey. They kill their prey by biting the throat, which damages the trachea and the major blood vessels of the neck.^[72] The size of the hunting pack is determined by the type of prey targeted, with large packs formed to help hunt large prey. Large prey can include kangaroos, cattle, water buffalo and wild horses.^[26] Dingoes will assess and target prey based on the prey's ability to inflict damage on dingoes. Large kangaroos are the most commonly killed prey. The main tactic is to sight the kangaroo, bail it up, then kill it. Dingoes typically hunt large kangaroos by having lead dingoes chase the quarry toward the paths of their packmates, which are skilled at cutting corners in chases. The kangaroo becomes exhausted and is then killed. This same tactic is used by wolves, African hunting dogs, and hyenas. Another tactic shared with African hunting dogs is a relay pursuit until the prey is exhausted. A pack of dingoes is three times as likely to bring down a kangaroo than an individual because the killing is done by those following the lead chaser, which has also become exhausted.^[25] There are two patterns of the final stage of the attack. An adult or juvenile kangaroo is nipped at the hamstrings of the hind legs to slow it before an attack to the throat. A small adult female or juvenile is bitten on the neck or back by dingoes running beside them.^[26] In one area of Central Australia, dingoes hunt kangaroos by chasing them into a wire fence where they become temporarily immobilised. The largest male red kangaroos tend to ignore dingoes, even when the dingoes are hunting the younger males and females. A large eastern grey kangaroo successfully fought off an attack by a single dingo which lasted for over an hour. Wallabies are hunted in a similar manner to kangaroos, the difference being that a single dingo will hunt using scent rather than sight and the hunt may last for several hours.^[25]

Dingo packs may attack young cattle and buffalo but never healthy, grown adults. They focus on the sick or injured young. The tactics include harassing a mother with young, panic a herd to separate the adults from the young, or watch a herd and look for any unusual behaviour that might then be exploited.^[25] One 1992 study in the Fortescue River region observed that cattle will defend their calves by circling around the calves or aggressively charging dingoes. In one study of 26 approaches, 24 were by more than one dingo and only 4 resulted in calves being killed. Dingoes often revisited carcasses. They did not touch fresh cattle carcasses until these were largely skin and bone, and even when these were plentiful they still preferred to hunt kangaroo. Out of 68 chases of sheep, 26 sheep were seriously injured but only 8 were killed. The dingoes could outrun the sheep and the sheep were defenceless. However, the dingoes in general appeared not to be motivated to kill sheep, and in many cases just loped alongside the sheep before veering off to chase another sheep. For those that did kill and consume sheep, there was still a large quantity of kangaroo in their diet, indicating once again a preference for kangaroo.^[73]

Lone dingoes can run down a rabbit, but are more successful by targeting kittens near rabbit warrens. Dingoes will take young nestlings or young birds, in addition to birds that are moulting and therefore cannot fly. In the coastal wetlands of northern Australia, dingoes depend on magpie geese for a large part of their diet and a lone dingo will sometimes distract these while a white breasted sea eagle makes a kill which is too heavy for it to carry off, with the dingo then driving the sea eagle away. They will also scavenge on prey dropped from the nesting platforms of sea eagles. Lone dingoes may hunt small rodents and grasshoppers in grass by using their sense of smell and hearing, then pouncing on them with their forepaws.^[25]

Competitors

Dingoes and their hybrids co-exist with the native Quoll. They also co-occur in the same territory as the introduced European red fox and feral cat, but little is known about the relationships between these three. Dingoes and their hybrids will drive off foxes from sources of water and occasionally eat feral cats. Dingoes can be killed by buffalo and cattle goring and kicking them, from snake bite, and predation on their pups by wedge-tailed eagles.^[26]

Barking

Dingo on the Nullarbor

Compared to most domestic dogs, the bark of a dingo is short and monosyllabic, and is rarely used. Barking was observed to make up only 5% of vocalisations. Dog barking has always been distinct from wolf barking.^[75] Australian dingoes bark mainly in swooshing noises or in a mixture of atonal and tonal sounds. In addition, barking is almost exclusively used for giving warnings. Warn-barking in a homotypical sequence and a kind of "warn-howling" in a heterotypical sequence have also been observed. The bark-howling starts with several barks and then fades into a rising and ebbing howl and is probably (similar to coughing) used to warn the puppies and members of the pack. Additionally, dingoes emit a sort of "wailing" sound, which they mostly use when approaching a wateringhole, probably to warn already present dingoes.^[76]

According to the present state of knowledge, it is not possible to get Australian dingoes to bark more frequently by putting them in contact with other domestic dogs. However, German zoologist Alfred Brehm reported a dingo that learned the more "typical" form of barking and how to use it, while its brother did not.^[77] Whether dingoes bark or bark-howl less frequently in general is not certain.^[74]

Howling

Dingoes have three basic forms of howling (moans, bark-howls and snuffs) with at least 10 variations. Usually, three kinds of howls are distinguished: long and persistent, rising and ebbing, and short and abrupt.

Observations have shown that each kind of howling has several variations, though their purpose is unknown. The frequency of howling varies with the season and time of day, and is also influenced by breeding, migration, lactation, social stability and dispersal behaviour. Howling can be more frequent in times of food shortage, because the dogs become more widely distributed within their home range.^[76]

Additionally, howling seems to have a group function, and is sometimes an expression of joy (for example, greeting-howls). Overall howling was observed less frequently in dingoes than among grey wolves. It may happen that one dog will begin to howl, and several or all other dogs will howl back and bark from time to time. In the wilderness, dingoes howl over long distances to attract other members of the pack, to find other dogs, or to keep intruders at bay. Dingoes howl in chorus with significant pitches, and with increasing number of pack-members, the variability of pitches also increases.^[78] Therefore, it is suspected that dingoes can measure the size of a pack without visual contact.^[79] Moreover, it has been proposed that their highly variable chorus howls may generate a confounding effect in the receivers by making pack size appear larger.^[80]

Other forms of communication

Growling, making up approximately 65% of the vocalisations, is used in an agonistic context for dominance, and as a defensive sound. Similar to many domestic dogs, a reactive usage of defensive growling is only rarely observed. Growling very often occurs in combination with other sounds, and has been observed almost exclusively in swooshing noises (similar to barking).^[74]

During observations in Germany, dingoes were heard to produce a sound that observers have called Schrappen. It was only observed in an agonistic context, mostly as a defence against obtrusive pups or for defending resources. It was described as a bite intention, during which the receiver is never touched or hurt. Only a clashing of the teeth could be heard.^[74]

Aside from vocal communication, dingoes communicate, like all domestic dogs, via scent marking specific objects (for example, Spinifex) or places (such as waters, trails and hunting grounds) using chemical signals from their urine, feces and scent glands. Males scent-mark more frequently than females, especially during the mating season. They also scent-rub, whereby a dog rolls its neck, shoulders, or back on something that is usually associated with food or the scent markings of other dogs.^[76]

Unlike wolves, dingoes can react to social cues and gestures from humans.^[81]

Social behaviour

The dingo's social behaviour is about as flexible as that of a coyote or gray wolf, which is perhaps one of the reasons it was initially believed that the dingo was descended from the Indian wolf.^[85] While young males are often solitary and nomadic in nature, breeding adults will often form a settled pack.^[86] However, in areas of the dingo's habitat with a widely spaced population, breeding pairs remain together, apart from others.^[86] Dingo distributions are a single dingo, 73%; two dingoes, 16%; three dingoes, 5%; four dingoes, 3%; and packs of five to seven dingoes, 3%. A dingo pack usually consists of a mated pair, their offspring from the current year and sometimes offspring from the previous year.^[27]

Where conditions are favourable among dingo packs, the pack is stable with a distinct territory and little overlap between neighbours.^[87] The size of packs often appears to correspond to the size of prey that appears in the pack's territory.^[87] Desert areas have smaller groups of dingoes with a more loose territorial behaviour and sharing of the water sites.^[88] It has been noted that the average monthly pack size was between three and twelve members.^[89]

Similar to other canids, a dingo pack largely consists of a mated pair, their current year's offspring, and occasionally a previous year's offspring.^[87] There are dominance hierarchies both between and within males and females, with males usually being more dominant than females.^[87] However, a few exceptions have been noted in captive packs.^[87] During travel, while eating prey, or when approaching a water source for the first time, the breeding male will be seen as the leader, or alpha.^[90] Subordinate dingoes will approach a more dominant dog in a slightly crouched posture, ears flat and tail down, to ensure peace in the pack.^[87] Establishment of artificial packs in captive dingoes has failed.^[87]

Reproduction

Dingo pups

Dingoes breed once annually, depending on the estrus cycle of the females which, according to most sources, only come in heat once per year. Dingo females can come in heat twice per year, but can only be pregnant once a year, with the second time only seeming to be pregnant.^[91][92]

Males are virile throughout the year in most regions, but have a lower sperm production during the summer in most cases. During studies on dingoes from the Eastern Highlands and Central Australia in captivity, no specific breeding cycle could be observed. All were potent throughout the year. The breeding was only regulated by the heat of the females. A rise in testosterone was observed in the males during the breeding season, but this was attributed to the heat of the females and copulation. In contrast to the captive dingoes, captured dingo males from Central Australia did show evidence of a male breeding cycle. Those dingoes showed no interest in females in heat (this time other domestic dogs) outside of the mating season (January to July) and did not breed with them.^[93]

The mating season usually occurs in Australia between March and May (according to other sources between April and June). During this time, dingoes may actively defend their territories using vocalisations, dominance behaviour, growling and barking.^[79]

Most females in the wild start breeding at the age of two years. Within packs, the alpha female tends to go into heat before subordinates and actively suppresses mating attempts by other females. Males become sexually mature between the ages of one and three years. The precise start of breeding varies depending on age, social status, geographic range and seasonal conditions. Among dingoes in captivity, the pre-estrus was observed to last 10–12 days. However, it is suspected that the pre-estrus may last as long as 60 days in the wild.^[76]

A male dingo with his pups

In general, the only dingoes in a pack that successfully breed are the alpha pair, and the other pack members help with raising the pups. Subordinates are actively prevented from breeding by the alpha pair and some subordinate females have a false pregnancy. Low-ranking or solitary dingoes can successfully breed if the pack structure breaks up.^[94]

The gestation period lasts for 61–69 days and the size of the litter can range from one to 10 (usually five) pups, with the number of males born tending to be higher than that of females. Pups of subordinate females usually get killed by the alpha female, which causes the population increase to be low even in good times. This behaviour possibly developed as an adaptation to the fluctuating environmental conditions in Australia. Pups are usually born between May and August (the winter period), but in tropical regions, breeding can occur at any time of the year.^[76]

At the age of three weeks, the pups leave the den for the first time, and leave it completely at eight weeks. In Australia, dens are mostly underground. There are reports of dens in abandoned rabbit burrows, rock formations, under boulders in dry creeks, under large spinifex, in hollow logs, in augmented burrows of monitor lizards and wombat burrows. The pups usually stray around the den within a radius of 3 km (2 mi), and are accompanied by older dogs during longer travels. The transition to consuming solid food is normally accompanied by all members of the pack during the age of 9 to 12 weeks. Apart from their own experiences, pups also learn through observation.^[95] Young dingoes usually become independent at the age of three to six months or they disperse at the age of 10 months when the next mating season starts.

Migration

Dingoes usually remain in one area and do not undergo seasonal migrations. However, during times of famine, even in normally "safe" areas, dingoes travel into pastoral areas, where intensive, human-induced control measures are undertaken. It was noted in Western Australia in the 1970s that young dogs can travel for long distances when necessary. About 10% of the dogs captured—all younger than 12 months—were later recaptured far away from their first location. Among these, 10% of the travelled distance for males was 21.7 km (13.5 mi) and for females 11 km (7 mi). Therefore, travelling dingoes had lower chances of survival in foreign territories, and it was apparently unlikely that they would survive long migrations through occupied territories. The rarity of long migration routes seemed to confirm this. During investigations in the Nullarbor Plain, even longer migration routes were recorded. The longest recorded migration route of a radio-collared dingo was about 24–32 km (15–20 mi).^[96]

Attacks on humans

Although dingoes are large enough to be dangerous, they generally avoid conflict with humans. Apart from the well-known case in which an infant was taken from a campsite (see below), there have been numerous confirmed dingo attacks, often involving people feeding wild dingoes, particularly on Fraser Island, a special center of dingo-related tourism (see main article). Most dingo attacks are minor in nature, but some can be major, and a few can be fatal. Many Australian national parks have signs advising visitors not to feed wildlife, partly because this practice is not healthy for the animals, and partly because it may encourage undesirable behaviour, such as snatching or biting by dingoes, kangaroos, goannas and some birds.

Extinctions

Some researchers propose that the dingo caused the extinction of the thylacine, the Tasmanian devil and the Tasmanian nativehen from mainland Australia because of the correlation in space and time with the dingo's arrival. Recent studies have questioned this proposal, suggesting that climate change and increasing human populations may have been the cause.^[97] Dingoes do not seem to have had the same ecological impact that the red fox had in later times. This might be connected to the dingo's way of hunting and the size of their favoured prey, as well as to the low number of dingoes in the time before European colonisation.^[98]

The assumption that dingoes and thylacines were competitors for the same prey stems from their external similarities; the thylacine had a stronger and more efficient bite, but was probably dependent on relatively small prey, while the dingo's stronger skull and neck would have allowed it to bring down bigger prey.^[99] The dingo was probably a superior hunter, as it hunted cooperatively in packs and could better defend resources, while the thylacine was probably more solitary. Also, wild dingo populations might have had demographic support from conspecific living with humans.

The extinction of the thylacine on the continent around 2,000 years ago has also been linked to changes in climate and land use by the Aborigines. It is plausible to name the dingo as the cause of the extinction, but significant morphological differences between the two suggest that the ecological overlapping of both species might be exaggerated. The dingo has the dentition of a generalist, while the thylacine had the dentition of a specialist carnivore without any signs of consumption of carrion or bones. It is also argued that the thylacine was a flexible predator that should have withstood the competition by the dingo, but was instead wiped out due to human persecution.

This theory does not explain how the Tasmanian devil and the dingo coexisted on the same continent until about 430 years ago, when the dingo supposedly caused the Tasmanian devil's demise. The group dynamics of dingoes should have successfully kept devils away from carrion, and since dingoes are able to break bones, little would have been left for the devils to scavenge. Additionally, devils are successful hunters of small- to medium-sized prey, so there should have been an overlapping of the species in this area, too. Furthermore, the arguments that the dingo caused the extinction of the thylacine, the devil and the hen are in direct conflict with each other. If the dingo were really so similar to the thylacine and the Tasmanian devil in its ecological role and suppressed both, then coexisting with both for such an extended time is strange. Although this is a possible result of the dingo's introduction, critics regard the evidence for this as insubstantial.^[100]

In 2017, a genetic study found that the population of the northwestern dingoes had commenced expanding since 4,000—6,000 years ago. This was proposed to be due to either their first arrival in Australia or due to the commencement of the extinction of the Tasmanian tiger with the dingo expanding into the Tasmanian tiger's former range.^[23]

Ecological impact

Dingo, Fraser Island, Queensland

The dingo is regarded as part of the native Australian fauna by many environmentalists and biologists, as these dogs existed on the continent before the arrival of the Europeans and a mutual adaptation of the dingoes and their surrounding ecosystems had occurred.

Much of the present place of wild dogs in the Australian ecosystem, especially in the urban areas, remains unknown. Although the ecological role of dingoes in Northern and Central Australia is well understood, the same does not apply to the role of wild dogs in the east of the continent. In contrast to some claims,^[101] dingoes are assumed to have a positive impact on biodiversity in areas where feral foxes are present.^[102]

Dingoes are regarded as apex predators and possibly perform an ecological key function. It is likely (with increasing evidence from scientific research) that they control the diversity of the ecosystem by limiting the number of prey and keeping the competition in check. Wild dogs hunt feral livestock such as goats and pigs, as well as native prey and introduced animals. The low number of feral goats in Northern Australia is possibly caused by the presence of the dingoes, but whether they control the goats' numbers or not is still disputable. Studies from 1995 in the northern wet forests of Australia found the dingoes there did not reduce the number of feral pigs, but their predation only affects the pig population together with the presence of water buffaloes (which hinder the pigs' access to food).^[103]

Observations concerning the mutual impact of dingoes and red fox and cat populations suggest dingoes limit the access of foxes and cats to certain resources. As a result, it is assumed that a disappearance of the dingoes may cause an increase of red fox and feral cat numbers and, therefore, a higher pressure on native animals. These studies found the presence of dingoes is one of the factors that keep fox numbers in an area low, and therefore reduces pressure on native animals, which then do not disappear from the area. The countrywide numbers of red foxes are especially high where dingo numbers are low, but other factors might responsible for this, depending on the area.^[104] Evidence was found for a competition between wild dogs and red foxes in the Blue Mountains of New South Wales, since there were many overlaps in the spectrum of preferred prey, but there was only evidence for local competition, not on a grand scale.^[105]

It is also possible that dingoes can live with red foxes and feral cats without reducing their numbers in areas with sufficient food resources (for example, high rabbit numbers) and hiding places. Nearly nothing is known about the relationship of wild dogs and feral cats, except both mostly live in the same areas. Although wild dogs also eat cats, it is not known whether this affects the cat populations.^[104] At the moment, the Invasive Animals Cooperative Research Centre is investigating the exact effects of dingoes on the fox and cat populations to determine the benefits of keeping the dog in certain areas of Australia.^[106] In many areas, wild dogs live together with the most species of quolls , except for the eastern quoll, which is probably extinct on the mainland, so wild dogs are not regarded as a threat to them.

Additionally, the disappearance of dingoes might increase the prevalence of kangaroo, rabbit and Australian brushturkey numbers. In the areas outside the Dingo Fence, the number of dingoes and emus is lower than in the areas inside. However, the numbers changed depending on the habitat. Since the environment is the same on both sides of the fence, the dingo was assumed to be a strong factor for the regulation of these species.^[107] Therefore, some people demand that dingo numbers should be allowed to increase or dingoes should be reintroduced in areas with low dingo populations to lower the pressure on endangered populations of native species and to reintroduce them in certain areas. In addition, the presence of the Australian brushturkey in Queensland increased significantly after dingo baiting was conducted.^[108]

Cultural impact

Cultural opinions about the dingo are often based on its perceived "cunning", and the idea that it is an intermediate between civilisation and wildness.^[109]

Some of the early European settlers looked on dingoes as domestic dogs, while others thought they were more like wolves. Over the years, dingoes began to attack sheep, and their relationship to the Europeans changed very quickly: they were regarded as devious and cowardly, since they did not fight bravely in the eyes of the Europeans, and vanished into the bush.^[110] Additionally, they were seen as promiscuous or as devils with a venomous bite or saliva, and so they could be killed unreservedly. Over the years, dingo trappers gained some prestige for their work, especially when they managed to kill hard to catch dingoes. Dingoes were associated with thieves, vagabonds, bushrangers and parliamentary opponents. From the 1960s, politicians began calling their opponents "dingo," meaning they were cowardly and treacherous, and it has become a popular form of attack since then.^[111] Today, the word "dingo" still stands for "coward" and "cheat," with verb and adjective forms used, as well.^[109]

The image of the dingo has ranged among some groups from the instructive^[112] to the demonic.^[113]

Ceremonies (like a keen at the Cape York Peninsula in the form of howling^[95]) and dreamtime stories are connected to the dingo, which were passed down through the generations.

The dingo plays a prominent role in the dreamtime stories of indigenous Australians,^[29] however it is rarely depicted in their cave paintings when compared with the extinct thylacine.^[30][14] One of the tribal elders of the people of the Yarralin, Northern Territory region tells that the dreamtime dingo is the ancestor of both dingoes and humans. The dingoes "are what we would be if we were not what we are."^[29]

Similar to how Europeans acquired dingoes, the Aboriginal people of Australia acquired dogs from the immigrants very quickly. This process was so fast that Francis Barrallier (surveyor on early expeditions around the colony at Port Jackson) discovered in 1802 that five dogs of European origin were there before him.^[111] One theory holds that other domestic dogs will adopt the role of the "pure" dingo.^[112] Introduced animals, such as the water buffalo and the domestic cat, have been adopted into the indigenous Aboriginal culture in the forms of rituals, traditional paintings and dreamtime stories.^[109]

Most of the published myths originate from the Western Desert and show a remarkable complexity. In some stories, dingoes are the central characters, while in others, they are only minor ones. One time, it is an ancestor from the dreamtime who created humans and dingoes or gave them their current shape. There are stories about creation, socially acceptable behaviour, and explanations why some things are the way they are. There are myths about shapeshifters (human to dingo or vice versa), "dingo-people," and the creation of certain landscapes or elements of those landscapes, like waterholes or mountains.

Economic impact

Livestock farming commenced expanding across Australia from the early 1800s, which led to conflict between the dingo and graziers. Sheep, and to a lesser extent cattle, are an easy target for dingoes. The pastoralists and the government bodies that support this industry have shot, trapped, and poisoned dingoes or destroyed dingo pups in their dens. After two centuries of persecution, the dingo or dingo–dog hybrids can still be found across most of the continent.^[28]

Research on the real extent of the damage and the reason for this problem only started recently. Livestock can die from many causes and, when the carcass is found, it is often difficult to determine with certainty the cause of death. Since the outcome of an attack on livestock depends to a high degree on the behaviour and experience of the predator and the prey, only direct observation is certain to determine whether an attack was by dingoes or another domestic dog. Even the existence of remnants of the prey in the scat of wild dogs do not prove they are pests, since wild dogs also eat carrion. Exact numbers or reliable estimates of the damage caused by wild dogs are, therefore, hard to obtain and are seldom reliable. Even if livestock is not a big part of the dingo's diet, the extent of damage dingoes could potentially cause to the livestock industry could be much larger because of wanton killing.

The significance of dingoes as a pest is based primarily on the predation of sheep and, to a lesser extent, on cattle, and is not connected only to the direct loss of livestock. Harassment of sheep can cause a less optimal use of grassland and miscarriages.

Distribution of wild dogs and livestock^[4]

The cattle industry can tolerate low to moderate, and sometimes high, grades of wild dogs (therefore dingoes are not so easily regarded as pests in these areas). In the case of sheep and goats, a zero-tolerance attitude is common. The biggest threats are dogs that live inside or near the paddock areas. The extent of sheep loss is hard to determine, due to the wide pasture lands in some parts of Australia. The numbers of cattle losses is much more variable and less well-documented.

In 2006, cattle losses in the Northern Territory rangeland grazing areas were estimated between 0-30%.^[94]

Therefore, factors such as availability of native prey, as well as the defending behaviour and health of the cattle, play an important role in the number of losses. A study in Central Australia in 2003 confirmed that dingoes only have a low impact on cattle numbers when a sufficient supply of other prey (such as kangaroos and rabbits) is available. In some parts of Australia, it is assumed that the loss of calves can be minimised if horned cattle are used instead of polled.^[79] The precise economic impact is not known in this case, and it is unlikely that the rescue of some calves compensates for the necessary costs of control measures. Calves usually suffer less lethal wounds than sheep due to their size and the protection by the adult cattle, and therefore have a higher chance of surviving an attack. As a result, the evidence of a dog attack may only be discovered after the cattle have been herded back into the enclosure, and signs such as bitten ears, tails and other wounds are discovered.

The opinions of cattle owners regarding dingoes are more variable than those of sheep owners. Some cattle owners believe that it is better that the weakened mother loses her calf in times of drought so that she does not have to care for her calf, too. Therefore, these owners are more hesitant to kill dingoes.^[95] The cattle industry may benefit from the predation of dingoes on rabbits, kangaroos and rats. Furthermore, the mortality rate of calves has many possible causes, and it is difficult to discriminate between them. The only reliable method to document the damage would be to document all pregnant cows, then observe their development and that of their calves.^[94] The loss of calves in observed areas where dingoes were controlled was higher than in other areas. Loss of livestock is, therefore, not necessarily caused by the occurrence of dingoes and is independent from wild dogs.^[114] One researcher has stated that for cattle stations where dingoes were controlled, kangaroos were abundant, and this affects the availability of grass.^[115]

Domestic dogs are the only terrestrial predators in Australia that are big enough to kill fully-grown sheep, and only a few sheep manage to recover from the severe injuries. In the case of lambs, death can have many causes apart from attacks by predators, which are blamed for the deaths because they eat from the carcasses. Although attacks by red foxes are possible, such attacks are more rare than previously thought.^[114] The fact that the sheep and goat industry is much more susceptible to damage caused by wild dogs than the cattle industry is mostly due to two factors: the flight behaviour of the sheep and their tendency to flock together in the face of danger, and the hunting methods of wild dogs, along with their efficient way of handling goat and sheep.

Therefore, the damage to the livestock industry does not correlate to the numbers of wild dogs in an area (except that there is no damage where no wild dogs occur^[114]).

According to a report from the government of Queensland, wild dogs cost the state about $30 million annually due to livestock losses, the spread of diseases and control measures. Losses for the livestock industry alone were estimated to be as high as $18 million.^[94] In Barcaldine, Queensland, up to one-fifth of all sheep are killed by dingoes annually, a situation which has been described as an "epidemic".^[116] According to a survey among cattle owners in 1995, performed by the Park and Wildlife Service, owners estimated their annual losses due to wild dogs (depending on the district) to be from 1.6% to 7.1%.^[117]

Despite the variety of estimates, there is little doubt that predation by dingoes can cause enormous economic damage, especially in times of drought when natural prey is sparse and the number of dingoes is still relatively high. Furthermore, wild dogs are involved in the spread of echinococcosis among cattle and sheep. An infection with echinococcosis can lead to confiscation of 90% of the intestines, which further leads to a value decrease of the meat and high economical damage.

Among the indigenous Australians, dingoes were also used as hunting aids, living hot water bottles and camp dogs. Their scalps were used as a kind of currency, their teeth were traditionally used for decorative purposes, and their fur for traditional costumes. In some parts of Australia, premiums are paid for dingo fur and scalps. The fur of dingoes generally has only a low value, and export of this fur is forbidden in states where they are protected. There is no widespread commercial catching and killing of dingoes for the purposes of obtaining their fur.

Sometimes "pure" dingoes are important for tourism, when they are used to attract visitors. However, this seems to be common only on Fraser Island, where the dingoes are extensively used as a symbol to enhance the attraction of the island. Tourists are drawn to the experience of personally interacting with dingoes. Pictures of dingoes appear on brochures, many websites and postcards advertising the island.^[118]

Deterrence

One method that does not have any proven effect is to hang dead dogs along the borders of the property in the belief that this would repel wild dogs.^[79]

Guardian animals

To protect livestock, livestock guardian dogs (for example, Maremmas), donkeys, alpacas and llamas are used.^[121][122]

To keep wild dogs away from certain areas, efforts are taken to make these areas unattractive for them (for example, by getting rid of food waste) and therefore forcing them to move elsewhere. Control through deliberately spreading disease is normally not considered. Such attempts probably would not be successful, because typical dog diseases are already present in the population. Additionally, dogs under human care would also be susceptible. Other biological control methods are not regarded as achievable, since there would be a high risk of decimating dogs under human care.

Dingo Fence

A part of the dingo fence

In the 1920s, the Dingo Fence was erected on the basis of the Wild Dog Act (1921) and, until 1931, thousands of miles of Dingo Fences had been erected in several areas of South Australia. In the year 1946, these efforts were directed to a single goal, and the Dingo Fence was finally completed. The fence connected with other fences in New South Wales and Queensland. The main responsibilities in maintaining the Dingo Fence still lies with the landowners, whose properties border on the fence and receive financial support from the government.

Reward system

A reward system (local, as well from the government) was active from 1846 to the end of the 20th century, but there is no evidence that – despite the billions of dollars spent – it was ever an efficient control method. Therefore, its importance declined over time.^[76]

Dingo scalping commenced in 1912 with the passage of the Wild Dogs Act by the government of South Australia. In an attempt to reduce depredation on livestock, that government offered a bounty for dingo skins, and this program was later repeated in Western Australia and the Northern Territory. One writer argues that this new legislation and economic driver had significant impacts on Aboriginal society in the region.^[123]

Poisoning

Warning of poisonous sodium fluoroacetate baits

Dead wild dogs that have been hung on a fence

Strychnine is still used in all parts of Australia.

Baits with the poison 1080 are regarded as the fastest and safest method for dog control, since they are extremely susceptible. Even small amounts of poison per dog are sufficient (0.3 mg per kg).^[94] The application of aerial baiting is regulated in the Commonwealth by the Civil Aviation Regulations (1988). The assumption that the tiger quoll might be damaged by the poison led to the dwindling of areas where aerial baiting could be performed. In areas where aerial baiting is no longer possible, it is necessary to put down baits.

Over the last years, cyanide-ejectors and protection collars (filled with 1080 on certain spots) have been tested.^[124][125]

The eradication of dingoes due to livestock damage decreased along with the importance of the sheep industry and the usage of strychnine (which beforehand had been used for 100 years) in the 1970s. The number of doggers also decreased and the frequency of government-approved aerial baiting increased. During this period, many farmers in Western Australia switched to the cattle industry, and findings in the area of biology led to a significant change in control measures and techniques in association with reduced costs and increased efficiency. At the same time, the importance of 1080 increased.

In 2016, controversy surrounded a plan to inject a population of dingoes on Pelorus Island, off the coast of northern Queensland, Australia, with pills that would release a fatal dose of 1080 poison two years after the dingoes were to be intentionally released to help eradicate goats. The dingoes were dubbed 'death-row dingoes', and the plan was blocked due to concerns for a locally threatened shorebird.^[126]

Neutering

Owners of dingoes and other domestic dogs are sometimes asked to neuter their pets and keep them under observation to reduce the number of stray/feral dogs and prevent interbreeding with dingoes.^[94]

Efficiency of measures

The efficiency of control measures was questioned in the past and is often questioned today, as well as whether they stand in a good cost-benefit ratio. The premium system proved to be susceptible to deception and to be useless on a large scale, and can therefore only be used for getting rid of "problem-dogs".^[79][127] Animal traps are considered inhumane and inefficient on a large scale, due to the limited efficacy of baits. Based on studies, it is assumed that only young dogs that would have died anyway can be captured.^[96] Furthermore, wild dogs are capable of learning and sometimes are able to detect and avoid traps quite efficiently. In one case, a dingo bitch followed a dogger and triggered his traps one after another by carefully pushing her paw through the sand that covered the trap.^[110]

Poisonous baits can be very effective when they are of good meat quality; however, they do not last long^[128] and are occasionally taken by red foxes, quolls, ants and birds. Aerial baiting can nearly eliminate whole dingo populations.^[96] Livestock guardian dogs can effectively minimise livestock losses, but are less effective on wide open areas with widely distributed livestock. Furthermore, they can be a danger to the livestock or be killed by control measures themselves when they are not sufficiently supervised by their owners.^[125] Fences are reliable in keeping wild dogs from entering certain areas, but they are expensive to build, need permanent maintenance, and only cause the problem to be relocated.

Control measures mostly result in smaller packs and a disruption of pack structure. The measures seem to be rather detrimental to the livestock industry because the empty territories are taken over by young dogs and the predation then increases. Nonetheless, it is regarded as unlikely that the control measures could completely eradicate the dingo in Central Australia, and the elimination of all wild dogs is not considered a realistic option.

It has been shown that culling a small percentage of immature dingoes on Fraser Island have little significant negative impact on the overall island population, though this is being disputed.^[129]

Hybridisation

Broad distribution map of dingoes and dingo-dog hybrids showing percent purity.^[137]

Although dingo-like, this wild dog has an atypical colouration and is therefore most likely a dingo-crossbreed.

European domestic dogs first arrived in Australia during the European colonisation. These dogs reverted to the wild (both unintentionally and intentionally), produced feral populations and interbred with the existing dingoes. Hybrids of dingoes and domestic dogs exist today in all wild dog populations of Australia, with their numbers having increased to such a degree that any completely "pure" populations may no longer exist.^[111]

Dingo-like domestic dogs and dingo-hybrids can be generally distinguished from "pure" dingoes by their fur colour, since there is a wider range of colours and patterns among them than among dingoes. In addition, the more dog-typical kind of barking exists among the hybrids, and differences in the breeding cycle,^[138] certain skull characteristics,^[139] and genetic analyses^[140] can be used for differentiation. Despite all the characteristics that can be used for distinguishing between dingoes and other domestic dogs, there are two problems that should not be underestimated. First, there is no real clarity regarding at what point a dog is regarded as a "pure" dingo,^[141] and, secondly, no distinguishing feature is completely reliable—it is not known which characteristics permanently remain under the conditions of natural selection.

There are two main opinions regarding this process of interbreeding. The first, and likely most common, position states that the "pure" dingo should be preserved via strong controls of the wild dog populations, and only "pure" or "nearly-pure" dingoes should be protected.^[142] The second position is relatively new and is of the opinion that people must accept that the dingo has changed and that it is impossible to bring the "pure" dingo back. Conservation of these dogs should therefore be based on where and how they live, as well as their cultural and ecological role, instead of concentrating on precise definitions or concerns about "genetic purity".^[143] Both positions are controversially discussed.

Due to this interbreeding, there is a wider range of fur colours, skull shapes and body size in the modern-day wild dog population than in the time before the arrival of the Europeans. Over the course of the last 40 years, there has been an increase of about 20% in the average wild dog body size.^[144] It is currently unknown whether, in the case of the disappearance of "pure" dingoes, remaining hybrids would alter the predation pressure on other animals. It is also unclear what kind of role these hybrids would play in the Australian ecosystems. However, it is unlikely that the dynamics of the various ecosystems will be excessively disturbed by this process.^[79]

In 2011, a total of 3,941 samples were included in the first continent-wide DNA study of wild dogs. The study found that 46% were pure dingoes which exhibited no dog alleles (gene expressions). There was evidence of hybridisation in every region sampled. In Central Australia only 13% were hybrids, however in southeastern Australia 99% were hybrids or feral dogs. Pure dingo distribution was 88% in the Northern Territory, intermediate numbers in Western Australia, South Australia and Queensland, and 1% in New South Wales and Victoria. Almost all wild dogs showed some dingo ancestry,^[145][146] with only 3% of dogs showing less than 80% dingo ancestry. This indicates that domestic dogs have a low survival rate in the wild or that most hybridisation is the result of roaming dogs that return to their owners. No populations of feral dogs have been found in Australia.^[145]

In 2016, a three dimensional geometric morphometric analysis of the skulls of dingoes, dogs and their hybrids found that dingo-dog hybrids exhibit morphology closer to the dingo than to the parent group dog. Hybridisation did not push the unique Canis dingo cranial morphology towards the wolf phenotype, therefore hybrids cannot be distinguished from dingoes based on cranial measures. The study suggests that the wild dingo morphology is dominant when compared with the recessive dog breed morphology, and concludes that although hybridisation introduces dog DNA into the dingo population, the native cranial morphology remains resistant to change.^[146]

Criticism

The ownership of dingoes as pets and their breeding is widely criticised. The main criticism is that the activities and the resulting consequences of the dingo conservation groups, "dingo farms" and legislation for legal ownership of dingoes for people in public, is seen to be an additional threat to the survival of the pure dingoes. This fear exists because the majority of these breeding activities effectively expedite the interbreeding of dingoes and other domestic dogs, when the identification of a pure dingo is not absolutely correct respectively when hybrids are sold as "pure" dingoes.^[76]

Supporters of breeding programmes are only mildly optimistic about a successful outcome. Success in the form of a population viable for future re-wilding cannot be easily accomplished.^[149] According to David Jenkins, the breeding and reintroduction of pure dingoes is no easy option and, at the time, there were no studies that seriously dealt with this topic, especially in areas where dingo populations are already present.^[150]

An additional threat is that breeders may unconsciously select tamer dingoes by breeding individuals who are easier to manage. Therefore, it may happen that, over the years, the tame populations may become less suitable for living in the wild than their ancestors. In addition, a loss of genetic diversity (thus resulting in a higher susceptibility to diseases) might occur due to a small founding population, and negative changes could occur simply because the dogs were captive-bred. Furthermore, some features that are necessary for survival in the wild might "fade" under the conditions of domestication (for example, hunting techniques) because they are no longer needed.

Dingo pets are likely to escape.^[151]