Dingo (taxon)

Canis lupus dingo
Australian dingo
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: Canis lupus
Subspecies: C. lupus dingo[2]
Trinomial name
Canis lupus dingo[3]
(Canis dingo Meyer, 1793)[4]
Synonyms[5]

Canis dingo (Meyer, 1793), antarticus (Kerr, 1792) [suppressed, ICZN O.451], Canis australasiae (Desmarest, 1820), Canis australiae (Gray, 1826), Canis dingoides (Matschie, 1915), Canis macdonnellensis (Matschie, 1915), Canis novaehollandiae (Voigt, 1831), Canis papuensis (Ramsay, 1879), Canis tenggerana (Kohlbrugge, 1896), Canis hallstromi (Troughton, 1957), Canis harappensis (Prashad, 1936)

The taxon dingo always refers to the native dog found in Australia, but may at times also refer to other dogs native to peninsular and island southeast Asia and neighboring regions, such as the New Guinea singing dog (halstromi). It has variously been considered a species of the dog genus (Canis dingo), a subspecies of gray wolf (Canis lupus dingo), and an invalid taxon or taxonomically synonymous with Canis lupus familiaris.

The genetic evidence indicates that the dingo clade originated from East Asian domestic dogs and was introduced through the South-East Asian archipelago into Australia,[6][7] with a common ancestry between the Australian dingo and the New Guinea Singing Dog.[7][8]

Taxonomy

In 1758, the taxonomist Carl Linnaeus published in Systema Naturae a categorization of species which included the Canis species. Canis is a Latin word meaning dog.[9] In 1978, a review aimed at reducing the number of recognized Canis species proposed that "Canis dingo is now generally regarded as a distinctive feral domestic dog. Canis familiaris is used for domestic dogs, although taxonomically it should probably be synonymous with Canis lupus."[10] In 1982, the first edition of Mammal Species of the World listed Canis familiaris under Canis lupus with the comment: "Probably ancestor of and conspecific with the domestic dog, familiaris. Canis familiaris has page priority over Canis lupus, but both were published simultaneously in Linnaeus (1758), and Canis lupus has been universally used for this species",[11] which avoided classifying the wolf as the family dog. The dingo was then regarded as a domestic dog.

In 2003, the ICZN ruled in its Opinion 2027 that if wild animals and their domesticated derivatives are regarded as one species, then the scientific name of that species is the scientific name of the wild animal. In 2005, the third edition of Mammal Species of the World upheld Opinion 2027 with the name Lupus and the note: "Includes the domestic dog as a subspecies, with the dingo provisionally separate - artificial variants created by domestication and selective breeding. Although this may stretch the subspecies concept, it retains the correct allocation of synonyms."[2] The dingo is now listed among the many other Latin-named subspecies of Canis lupus as Canis lupus dingo.[3]

New Guinea Singing Dog

Taxonomic synonyms

Current taxonomy identifies 10 taxonomic synonyms for the term Canis lupus dingo, the majority of which referred to the Australian dingo, one to the New Guinea Singing Dog, and 3 that refer to dogs that were once thought to be dingoes that were found in Southern and Southeast Asia.

"dingo", Australian dingo

Included Canis dingo, antarticus [suppressed], Canis australasiae, Canis australiae, Canis dingoides, Canis macdonnellensis, Canis novaehollandiae.[12]

Main article: Dingo

"antarticus", [suppressed]

The name Canis antarticus (Kerr, 1792 – note: spelling is not antarcticus) was the first name designated but it was overlooked in preference to dingo. In 1947, the name was proposed for resurrection but was subsequently suppressed by Opinion 451 of the ICZN (1957A:331) in favour of Canis dingo.[13]

"hallstromi", New Guinea singing dog

"harappensis", ancient dog found in South Asia

During excavation seasons in 1924-25 and 1930–31, a team of researchers at Harappa, in modern Pakistan, collected a wide variety of ancient domestic animal remains which had been buried since three thousand years BC. The remains included a dog which the researcher who wrote the report named the Canis tenggeranas harappensis, noting "marked skull affinities to the Indian wolf" and hypothesized to be the ancestor of the Indian Greyhound.[14]

"tenggerana", Java

In the late nineteenth century, naturalists working in the Tennger Mountains in Eastern Java identified a dog which they named "Canis familiaris tenggerana" after the local people called the Tenggerese.[15] A similar dog existed in the Dieng highlands, and it is assumed that pure populations of these two dogs no longer exist, mainly as a result of cross-breeding with varieties of domestic dogs.[16]

"papuensis", Papua New Guinea

In a report to the Linnean Society of New South Wales in 1882, N. De Miklouho-Maclay identified several anatomical and behavioural differences between Australian Dingoes and dogs that inhabited the coastal lowlands (Maclay Coast) of Papua New Guinea. He gave the Papuan coastal canines the scientific name Canis papuensis. The differences between Canis dingo and Canis papuensis included a much smaller brain, which he attributed to the quite different lifestyles of the two animals. There were several differences in overall looks and build. Whereas the Australian Dingo is well known for its intelligence and boldness, as well as near independence from humans, he reported that the coastal Papuan canines remained on the periphery of native villages, regularly feeding on cast-offs and human waste. Hunting on their own was almost unknown. Instead of the bold independence of the Australian dingo, the coastal dogs behaved very subserviently toward humans, exhibiting begging and grovelling. Additionally, he stated, "The Canis papuensis is very different in appearance and character from Canis dingo; is generally smaller, has not the bushy tail of the dingo...." [17]

Taxonomic challenge

The taxonomic classification Canis lupus dingo has been challenged regarding both the New Guinea Singing Dog[18][19][20][21] and the Australian Dingo.[22] The argument is that these are distinctive wild forms that differ from wolves by behavioral, morphological and molecular characteristics, and therefore are effectively a diagnosable species, isolated in undisturbed natural environments, and thus can be considered a distinct taxon Canis dingo. This argument has been counter-challenged in that it does not duly recognise the relationship with Canis familiaris - they overlap morphologically, hybridize readily, and form a genetic continuum with dogs.[12]

Lineage

Ancestors

The Sahul Shelf and the Sunda Shelf during the past 12,000 years. Tasmania separated from the mainland 12,000 YBP,[23] New Guinea separated from the mainland 8,500 YBP.[24]

In 1995, a researcher compared the skull morphology of the dingo to those of other dogs and wolves and concluded that the dingo was a primitive dog that had evolved from an Asian wolf.[25]

In 2004, a study looked at the mtDNA control region (maternal mDNA) of the Australian dingo. All dingo sequences in the study were found to be either identical to, or differing by a single substitution from, a single mDNA haplotype named A29.[6] All dingo sequences since studied exhibit haplotype A29, which falls within the Clade A haplogroup that represents 70% of domestic dogs.[7][26][27]

In 2009, a study further categorised haplotype A29 within the a2 sub-haplogroup that originated from hybridization between dog and wolf post-domestication.[26][28] Later studies analysed the whole genome of a dingo provided significant evidence of admixture between the dingo and a Chinese wolf.[29][30] The dingo lineage derives from dog/Chinese wolf hybridization or genetic admixture.

Haplotype A29 is found in both Australian dingoes and in domestic dogs exclusively in the East Asian region: East Siberia, Arctic America, Japan, Indonesia, New Guinea[6] and in South China, Kalimantan, and Bali.[7] The evidence suggests that the haplotype was introduced from East Asia through the islands of the South-East Asian archipelago and into Australia.[6][7] Haplotype A29 was one of several domestic dog mDNA haplotypes brought into island South-East Asia but only A29 reached mainland Australia.[6]

At the end of the Last glacial maximum and the rise in sea levels, Tasmania became separated from the Australian mainland 12,000 years before present (YBP),[23] and New Guinea 8,500 YBP[24] by the inundation of the Sahul Shelf.[31] The oldest Canis lupus dingo remains found date back to 3,450 YBP from the Nullabor plain in the southern part of the Australian continent, and there are doubts about 8,000 YBP for other remains due to problems with their stratigraphic interpretation.[32] No remains have been uncovered in Tasmania, therefore the Dingo is estimated to have arrived in Australia between 3,500-12,000 YBP.[6] The timing suggested by mDNA genetic divergence is through a single founding event[6] or no more than a few founding events[8] either 4,600-5,400 YBP or 4,600-10,800 YPB,[6] or 4,640-18,300 YBP,[7] depending on mutation rate assumptions. They remained isolated from other dogs until the arrival of Europeans.[6][8]

Sister

In 2011, a study looked at the mtDNA control region (maternal mDNA) of the Australian dingo and the New Guinea Singing Dog. The mDNA haplotype A29, or a haplotype one step away, was found in all of the Australian dingoes and New Guinea Singing Dogs studied, indicating a common ancestry.[7]

In 2012, a study looked at 14,437 base-pairs of Y chromosome DNA (paternal yDNA) of the Australian dingo, and found 2 yDNA haplotypes. Haplotype H3 could be found in domestic dogs in East Asia and Northern Europe. Haplotype H60 had not been previously reported but was one step away from haplotype H5 that could be found in East Asian domestic dogs. Only the New Guinea Singing Dog and dingoes from north-eastern Australia showed haplotype H60, which implies a genetic relationship and that the dingo reached Australia from New Guinea. Haplotype H60 and H3 could be found among the southern Australian dingoes with H3 dominant, but haplotype H3 could only be found in the west of the continent and may represent a separate entry from the northwest.[8] Earlier studies using other genetic markers had found the indigenous Bali dog more closely aligned with the Australian dingo than to those European and Asian breeds selected for comparison which indicated that the Bali dog was genetically diverse with a diverse history,[33][34][35] however only one percent exhibited the A29 mDNA haplotype.[7]

In 2012, a study of genome-wide SNP data from Indigenous Australians of the Northern Territory and people from southern India found evidence of a gene flow occurring 4,230 YBP. This is also approximately when changes in tool technology, food processing, and the dingo appear in the Australian archaeological record, suggesting that these may be related to the migration from India.[36]

Cousins

mDNA (maternal)

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 Arc2 to modern Polynesian, Indonesian and ancient New Zealand dogs, and the ancient Polynesian Arc1 is indistinguishable from a number of widespread modern haplotypes (K.Greig et al. 2015).[37]

In 2011, a study looked at 582 mDNA base-pairs of the mtDNA control region of dogs from Island South-East Asia and found that the two most common dog haplotypes of the Indonesian region, in particular Bali and Kalimantan, was mDNA haplotype A75 (40%) and "the dingo founder haplotype" A29 (8%). Also present were haplotypes A120 and A145.[7] All 4 haplotypes fall within the a2 mDNA sub-haplogroup.[26][28] The study also looked at archaeological specimens of ancient Polynesian dogs from which only a " short-haplotype" (263 mDNA base pairs - nucleotide positions 15,458-15,720) could be derived. This short haplotype was named Arc2 and corresponds to mDNA haplotypes A75 and A120, and it could be found in 70% of samples found as far away as Hawaii and New Zealand. No dogs from Taiwan nor the Philippines carried the dingo or Polynesian haplotypes, which indicates that dogs did not enter the Pacific from a north-eastern route.[7]

In 2015, a study looked at the mtDNA control region taken from ancient dog samples discovered at an archaeological site at Wairau Bar, New Zealand and found that the sequences corresponded to the a2 mDNA sub-haplogroup. The dog samples all carried the mDNA haplotype A192, which has only been reported in some modern village dogs from Bali, Indonesia. When a shorter sequence was compared to the ancient Polynesian haplotype Arc2, all of the Wairau Bar dogs matched Arc2. Dogs from Wairau Bar likely represent part of the initial population of dogs introduced to New Zealand, having arrived with people around the beginning of the fourteenth century.[37]

All of these dogs carry haplotypes that fall under the mDNA a2 sub-haplogroup and are therefore descendents of a dog/Chinese wolf hybrid ancestor. In 2015 the most comprehensive study of mDNA haplotypes to date found that the a2 sub-haplogroup represents 3% of all dogs in South East Asia, 22% in the Indian subcontinent and 16% in East Asia.[28]

yDNA (paternal)
In 2011, a study conducted a Y chromosome analysis of 338 Australian dingoes, New Guinea Singing Dogs, and village dogs from Island Southeast Asia. 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 Island Southeast Asia but it is common in Taiwan. One H5 specimen from Taiwan clustered with one H60 from Australia with the indication of a common 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 Island Southeast Asia, would reflect the last vestiges of the earlier ancient breeds.[38]

References

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  29. ↑ Freedman, Adam H.; Gronau, Ilan; Schweizer, Rena M.; Ortega-Del Vecchyo, Diego; Han, Eunjung; Silva, Pedro M.; Galaverni, Marco; Fan, Zhenxin; Marx, Peter; Lorente-Galdos, Belen; Beale, Holly; Ramirez, Oscar; Hormozdiari, Farhad; Alkan, Can; Vilà, Carles; Squire, Kevin; Geffen, Eli; Kusak, Josip; Boyko, Adam R.; Parker, Heidi G.; Lee, Clarence; Tadigotla, Vasisht; Siepel, Adam; Bustamante, Carlos D.; Harkins, Timothy T.; Nelson, Stanley F.; Ostrander, Elaine A.; Marques-Bonet, Tomas; Wayne, Robert K.; Novembre, John (16 January 2014). "Genome Sequencing Highlights Genes Under Selection and the Dynamic Early History of Dogs". PLOS Genetics. PLOS Org. 10 (1): e1004016. doi:10.1371/journal.pgen.1004016. PMC 3894170Freely accessible. PMID 24453982. Retrieved December 8, 2014.
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  33. ↑ Irion, Dawn N; Schaffer, Alison L; Grant, Sherry; Wilton, Alan N; Pedersen, Niels C (2005). "Genetic variation analysis of the Bali street dog using microsatellites". BMC Genetics. 6: 6. doi:10.1186/1471-2156-6-6. PMC 549630Freely accessible. PMID 15701179.
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  37. 1 2 Greig, Karen; Boocock, James; Prost, Stefan; Horsburgh, K. Ann; Jacomb, Chris; Walter, Richard; Matisoo-Smith, Elizabeth (2015). "Complete Mitochondrial Genomes of New Zealand's First Dogs". PLoS ONE. 10 (10): e0138536. doi:10.1371/journal.pone.0138536. PMC 4596854Freely accessible. PMID 26444283.
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