Taxonomic tangles posed by human association – the urgent need for an evidence-based review of dingo and domestic dog taxonomy and nomenclature

Kylie M. Cairns; Mike Letnic; Euan G. Ritchie; Justin W. Adams; Mathew S. Crowther; Christopher R. Dickman; Melanie Fillios; Jack Pascoe; Bradley P. Smith; Thomas M. Newsome

doi:10.1071/AM24052

REVIEW (Open Access)

Previous Contents Vol 47(3)

Taxonomic tangles posed by human association – the urgent need for an evidence-based review of dingo and domestic dog taxonomy and nomenclature

Kylie M. Cairns

^A ^B ^* , Mike Letnic ^A ^B , Euan G. Ritchie ^C , Justin W. Adams ^D ^E ^F , Mathew S. Crowther

^G , Christopher R. Dickman

^G , Melanie Fillios ^H , Jack Pascoe ^I , Bradley P. Smith

^J and Thomas M. Newsome ^G

+ Author Affiliations

- Author Affiliations

^A Evolution & Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW, Australia.

^B Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW, Australia.

^C School of Life and Environmental Sciences, Deakin University, Burwood, Vic, Australia.

^D Department of Anatomy and Developmental Biology, Biomedicine Discovery Institute, Monash University, Melbourne, Vic, Australia.

^E Geosciences, Museums Victoria, Melbourne, Vic, Australia.

^F Palaeo-Research Institute, Faculty of Humanities, University of Johannesburg, Johannesburg, South Africa.

^G School of Life and Environmental Sciences, The University of Sydney, Sydney, NSW, Australia.

^H Department of Archaeology and Palaeoanthropology, School of Humanities, Arts and Social Sciences, The University of New England, Armidale, NSW, Australia.

^I Quantitative and Applied Ecology Group, School of Food, Agriculture and Ecosystem Sciences, The University of Melbourne, Parkville, Vic, Australia.

^J College of Psychology, School of Human Health and Social Sciences, Central Queensland University Australia, Wayville, SA, Australia.

^* Correspondence to: k.cairns@unsw.edu.au

Handling Editor: Ross Goldingay

Australian Mammalogy 47, AM24052 https://doi.org/10.1071/AM24052

Submitted: 5 December 2024 Accepted: 26 July 2025 Published: 21 August 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing on behalf of the Australian Mammal Society. This is an open access article distributed under the Creative Commons Attribution-NonCommercial 4.0 International License (CC BY-NC)

Abstract

A serious challenge in modern taxonomy is whether, or how, relationships with humans have altered the evolutionary trajectory of species and how this should be dealt with in taxonomic nomenclature. Despite their historic treatment as a discrete taxon, the taxonomy and nomenclature of dingoes has been robustly debated for decades, in part due to their close evolutionary relationship to wolves and domestic dogs, as well as uncertainty about the role of humans in shaping their origins. One outcome of this ongoing scientific debate is a set of criteria regarding reproductive isolation, evolutionary distinctiveness and evolutionary origins that can be used to refine discussion about the appropriate taxonomy of dingoes. We review significant scientific advances in our knowledge of the distinct identity of dingoes in light of these criteria and discuss the need for scientists to assess the multi-stage process of domestication when considering species that may have interacted with humans over evolutionary timescales. Current evidence indicates that dingoes are on a different evolutionary trajectory to domestic dogs and thus we argue that the appropriate nomenclature for dingoes is Canis dingo or Canis lupus dingo, not Canis familiaris. We suggest that an independent evidence-based scientific review with expert consultation is one pathway through which taxonomic debate about dingoes and other controversial species could be resolved.

Keywords: Canis dingo, Canis familiaris, Canis lupus dingo, culturally significant species/entity, dingo, domestic dog, taxonomic nomenclature.

References

Australian Bureau of Statistics (2020). Regional Population, 2018–19. (ABS: Canberra.)

Ahlgren, H., Norén, K., Angerbjörn, A., and Lidén, K. (2016). Multiple prehistoric introductions of the mountain hare (Lepus timidus) on a remote island, as revealed by ancient DNA. Journal of Biogeography 43(9), 1786-1796.
| Crossref | Google Scholar |

Ahn, B., Kang, M., Jeon, H., Kim, J.-S., Jiang, H., Ha, J., and Park, C. (2023). Origin and population structure of native dog breeds in the Korean peninsula and East Asia. iScience 26, 106982.
| Crossref | Google Scholar | PubMed |

Allen, B. L., Allen, L. R., Ballard, G., Jackson, S. M., and Fleming, P. J. S. (2017). A roadmap to meaningful dingo conservation. Canid Biology and Conservation 20(11), 45-56.
| Google Scholar |

Animal Medicines Australia (2019). Pets in Australia: A national survey of pets and people, Animal Medicines Australia. Available at https://animalmedicinesaustralia.org.au/resources/pets-in-australia-a-national-survey-of-pets-and-people

Ardalan, A., Oskarsson, M., Natanaelsson, C., Wilton, A. N., Ahmadian, A., and Savolainen, P. (2012). Narrow genetic basis for the Australian dingo confirmed through analysis of paternal ancestry. Genetica 140, 65-73.
| Crossref | Google Scholar | PubMed |

Arendt, M., Cairns, K. M., Ballard, J. W. O., Savolainen, P., and Axelsson, E. (2016). Diet adaptation in dog reflects spread of prehistoric agriculture. Heredity 117, 301-306.
| Crossref | Google Scholar | PubMed |

Axelsson, E., Ratnakumar, A., Arendt, M.-L., Maqbool, K., Webster, M. T., Perloski, M., Liberg, O., Arnemo, J. M., Hedhammar, A., and Lindblad-Toh, K. (2013). The genomic signature of dog domestication reveals adaptation to a starch-rich diet. Nature 495(7441), 360-364.
| Crossref | Google Scholar | PubMed |

Ballard, J. W. O., and Wilson, L. A. B. (2019). The Australian dingo: untamed or feral? Frontiers in Zoology 16, 2.
| Crossref | Google Scholar | PubMed |

Balme, J., O’Connor, S., and Fallon, S. (2018). New dates on dingo bones from Madura Cave provide oldest firm evidence for arrival of the species in Australia. Scientific Reports 8(1), 9933.
| Crossref | Google Scholar | PubMed |

Bergström, A., Frantz, L., Schmidt, R., Ersmark, E., Lebrasseur, O., Girdland-Flink, L., Lin, A. T., Storå, J., Sjögren, K.-G., Anthony, D., Antipina, E., Amiri, S., Bar-Oz, G., Bazaliiskii, V. I., Bulatović, J., Brown, D., Carmagnini, A., Davy, T., Fedorov, S., Fiore, I., Fulton, D., Germonpré, M., Haile, J., Irving-Pease, E. K., Jamieson, A., Janssens, L., Kirillova, I., Horwitz, L. K., Kuzmanovic-Cvetković, J., Kuzmin, Y., Losey, R. J., Dizdar, D. L., Mashkour, M., Novak, M., Onar, V., Orton, D., Pasarić, M., Radivojević, M., Rajković, D., Roberts, B., Ryan, H., Sablin, M., Shidlovskiy, F., Stojanović, I., Tagliacozzo, A., Trantalidou, K., Ullén, I., Villaluenga, A., Wapnish, P., Dobney, K., Götherström, A., Linderholm, A., Dalén, L., Pinhasi, R., Larson, G., and Skoglund, P. (2020). Origins and genetic legacy of prehistoric dogs. Science 370(6516), 557.
| Crossref | Google Scholar | PubMed |

Bergström, A., Stanton, D. W. G., Taron, U. H., Frantz, L., Sinding, M. S., Ersmark, E., Pfrengle, S., Cassatt-Johnstone, M., Lebrasseur, O., Girdland-Flink, L., Fernandes, D. M., Ollivier, M., Speidel, L., Gopalakrishnan, S., Westbury, M. V., Ramos-Madrigal, J., Feuerborn, T. R., Reiter, E., Gretzinger, J., Münzel, S. C., Swali, P., Conard, N. J., Carøe, C., Haile, J., Linderholm, A., Androsov, S., Barnes, I., Baumann, C., Benecke, N., Bocherens, H., Brace, S., Carden, R. F., Drucker, D. G., Fedorov, S., Gasparik, M., Germonpré, M., Grigoriev, S., Groves, P., Hertwig, S. T., Ivanova, V. V., Janssens, L., Jennings, R. P., Kasparov, A. K., Kirillova, I. V., Kurmaniyazov, I., Kuzmin, Y. V., Kosintsev, P. A., Lázničková-Galetová, M., Leduc, C., Nikolskiy, P., Nussbaumer, M., O’Drisceoil, C., Orlando, L., Outram, A., Pavlova, E. Y., Perri, A. R., Pilot, M., Pitulko, V. V., Plotnikov, V. V., Protopopov, A. V., Rehazek, A., Sablin, M., Seguin-Orlando, A., Storå, J., Verjux, C., Zaibert, V. F., Zazula, G., Crombé, P., Hansen, A. J., Willerslev, E., Leonard, J. A., Götherström, A., Pinhasi, R., Schuenemann, V. J., Hofreiter, M., Gilbert, M. T. P., Shapiro, B., Larson, G., Krause, J., Dalén, L., and Skoglund, P. (2022). Grey wolf genomic history reveals a dual ancestry of dogs. Nature 607(7918), 313-320.
| Crossref | Google Scholar | PubMed |

Botigué, L. R., Song, S., Scheu, A., Gopalan, S., Pendleton, A. L., Oetjens, M., Taravella, A. M., Seregély, T., Zeeb-Lanz, A., Arbogast, R.-M., Bobo, D., Daly, K., Unterländer, M., Burger, J., Kidd, J. M., and Veeramah, K. R. (2017). Ancient European dog genomes reveal continuity since the Early Neolithic. Nature Communications 8(1), 16082.
| Crossref | Google Scholar | PubMed |

Brown, S. K., Darwent, C. M., and Sacks, B. N. (2013). Ancient DNA evidence for genetic continuity in Arctic dogs. Journal of Archaeological Science 40(2), 1279-1288.
| Crossref | Google Scholar |

Cagan, A., and Blass, T. (2016). Identification of genomic variants putatively targeted by selection during dog domestication. BMC Ecology and Evolution 16, 10.
| Crossref | Google Scholar | PubMed |

Cairns, K. M. (2021). What is a dingo – origins, hybridisation and identity. Australian Zoologist 41(3), 322-337.
| Crossref | Google Scholar |

Cairns, K. M., and Wilton, A. N. (2016). New insights on the history of canids in Oceania based on mitochondrial and nuclear data. Genetica 144(5), 553-565.
| Crossref | Google Scholar | PubMed |

Cairns, K. M., Shannon, L. M., Koler-Matznick, J., Ballard, J. W. O., and Boyko, A. R. (2018). Elucidating biogeographical patterns in Australian native canids using genome wide SNPs. PLoS One 13(6), e0198754.
| Crossref | Google Scholar | PubMed |

Cairns, K. M., Crowther, M. S., Nesbitt, B., and Letnic, M. (2021). The myth of wild dogs in Australia: are there any out there? Australian Mammalogy 44, 67-75.
| Crossref | Google Scholar |

Cairns, K. M., Crowther, M. S., Parker, H. G., Ostrander, E. A., and Letnic, M. (2023). Genome-wide variant analyses reveal new patterns of admixture and population structure in Australian dingoes. Molecular Ecology 32, 4133-4150.
| Crossref | Google Scholar | PubMed |

Clutton-Brock, J. (1992). The process of domestication. Mammal Review 22(2), 79-85.
| Crossref | Google Scholar |

Clutton-Brock, J. (2015). Naming the scale of nature. In ‘Taxonomic Tapestries’. Eds A. M. Behie, M. F. Oxenham.) pp. 171–182. (ANU Press.)

Coates, D. J., Byrne, M., and Moritz, C. (2018). Genetic Diversity and Conservation Units: Dealing With the Species-Population Continuum in the Age of Genomics. Frontiers in Ecology and Evolution 6, 165.
| Crossref | Google Scholar |

Corbett, L. K. (2001). ‘The Dingo in Australia and Asia’. (University of NSW Press: Sydney)

Crowther, M. S., Fillios, M., Colman, N., and Letnic, M. (2014). An updated description of the Australian dingo (Canis dingo Meyer, 1793). Journal of Zoology 293, 192-203.
| Crossref | Google Scholar |

Cursino, M. S., Harriott, L., Allen, B. L., Gentle, M., and Leung, L. K. P. (2017). Do female dingo–dog hybrids breed like dingoes or dogs? Australian Journal of Zoology 65(2), 112-119.
| Google Scholar |

Daniels, M. J., and Corbett, L. (2003). Redefining introgressed protected mammals: when is a wildcat a wild cat and a dingo a wild dog? Wildlife Research 30(3), 213-218.
| Crossref | Google Scholar |

De Queiroz, K. (2007). Species Concepts and Species Delimitation. Systematic Biology 56(6), 879-886.
| Crossref | Google Scholar | PubMed |

Driscoll, C. A., Macdonald, D. W., and O’brien, S. J. (2009). From wild animals to domestic pets, an evolutionary view of domestication. Proceedings of the National Academy of Sciences of the United States of America 106 Suppl 1(Supplement 1), 9971-8.
| Crossref | Google Scholar | PubMed |

Druzhkova, A. S., Thalmann, O., Trifonov, V. A., Leonard, J. A., Vorobieva, N. V., Ovodov, N. D., Graphodatsky, A. S., and Wayne, R. K. (2013). Ancient DNA analysis affirms the canid from Altai as a primitive dog. PLoS One 8(3), e57754.
| Crossref | Google Scholar | PubMed |

Fan, Z., Silva, P., Gronau, I., Wang, S., Armero, A. S., Schweizer, R. M., Ramirez, O., Pollinger, J., Galaverni, M., Ortega Del-Vecchyo, D., Du, L., Zhang, W., Zhang, Z., Xing, J., Vilà, C., Marques-Bonet, T., Godinho, R., Yue, B., and Wayne, R. K. (2016). Worldwide patterns of genomic variation and admixture in gray wolves. Genome Research 26, 163-73.
| Crossref | Google Scholar | PubMed |

Feuerborn, T. R., Carmagnini, A., Losey, R. J., Nomokonova, T., Askeyev, A., Askeyev, I., Askeyev, O., Antipina, E. E., Appelt, M., Bachura, O. P., Beglane, F., Bradley, D. G., Daly, K. G., Gopalakrishnan, S., Murphy Gregersen, K., Guo, C., Gusev, A. V., Jones, C., Kosintsev, P. A., Kuzmin, Y. V., Mattiangeli, V., Perri, A. R., Plekhanov, A. V., Ramos-Madrigal, J., Schmidt, A. L., Shaymuratova, D., Smith, O., Yavorskaya, L. V., Zhang, G., Willerslev, E., Meldgaard, M., Gilbert, M. T. P., Larson, G., Dalén, L., Hansen, A. J., Sinding, M.-H. S., and Frantz, L. (2021). Modern Siberian dog ancestry was shaped by several thousand years of Eurasian-wide trade and human dispersal. Proceedings of the National Academy of Sciences of the United States of America 118(39), e2100338118.
| Crossref | Google Scholar | PubMed |

Field, M. A., Yadav, S., Dudchenko, O., Esvaran, M., Rosen, B. D., Skvortsova, K., Edwards, R. J., Keilwagen, J., Cochran, B. J., Manandhar, B., Bustamante, S., Rasmussen, J. A., Melvin, R. G., Chernoff, B., Omer, A., Colaric, Z., Chan, E. K. F., Minoche, A. E., Smith, T. P. L., Gilbert, M. T. P., Bogdanovic, O., Zammit, R. A., Thomas, T., Aiden, E. L., and Ballard, J. W. O. (2022). The Australian dingo is an early offshoot of modern breed dogs. Science Advances 8(16), eabm5944.
| Crossref | Google Scholar | PubMed |

Fillios, M. A., and Taçon, P. S. C. (2016). Who let the dogs in? A review of the recent genetic evidence for the introduction of the dingo to Australia and implications for the movement of people. Journal of Archaeological Science: Reports 7, 782-792.
| Crossref | Google Scholar |

Fogg, B. R., Howe, N., and Pierotti, R. (2015). Relationships between Indigenous American Peoples and Wolves 1: Wolves as Teachers and Guides. Journal of Ethnobiology 35(2), 262-285.
| Crossref | Google Scholar |

Frantz, L. A. F., Mullin, V. E., Pionnier-Capitan, M., Lebrasseur, O., Ollivier, M., Perri, A., Linderholm, A., Mattiangeli, V., Teasdale, M. D., Dimopoulos, E. A., Tresset, A., Duffraisse, M., McCormick, F., Bartosiewicz, L., Gál, E., Nyerges, É. A., Sablin, M. V., Bréhard, S., Mashkour, M., Bălăşescu, A., Gillet, B., Hughes, S., Chassaing, O., Hitte, C., Vigne, J.-D., Dobney, K., Hänni, C., Bradley, D. G., and Larson, G. (2016). Genomic and archaeological evidence suggest a dual origin of domestic dogs. Science 352(6290), 1228-1231.
| Crossref | Google Scholar | PubMed |

Freedman, A. H., and Wayne, R. K. (2017). Deciphering the Origin of Dogs: From Fossils to Genomes. Annual Review of Animal Biosciences 5, 281-307.
| Crossref | Google Scholar | PubMed |

Freedman, A. H., Gronau, I., Schweizer, R. M., Ortega-Del Vecchyo, D., Han, E., Silva, P. M., Galaverni, M., Fan, Z., Marx, P., Lorente-Galdos, B., Beale, H., Ramirez, O., Hormozdiari, F., Alkan, C., Vilà, C., Squire, K., Geffen, E., Kusak, J., Boyko, A. R., Parker, H. G., Lee, C., Tadigotla, V., Siepel, A., Bustamante, C. D., Harkins, T. T., Nelson, S. F., Ostrander, E. A., Marques-Bonet, T., Wayne, R. K., and Novembre, J. (2014). Genome Sequencing Highlights the Dynamic Early History of Dogs. PLoS Genetics 10(1), e1004016.
| Crossref | Google Scholar | PubMed |

Freedman, A. H., Schweizer, R. M., Ortega-Del Vecchyo, D., Han, E., Davis, B. W., Gronau, I., Silva, P. M., Galaverni, M., Fan, Z., Marx, P., Lorente-Galdos, B., Ramirez, O., Hormozdiari, F., Alkan, C., Vilà, C., Squire, K., Geffen, E., Kusak, J., Boyko, A. R., Parker, H. G., Lee, C., Tadigotla, V., Siepel, A., Bustamante, C. D., Harkins, T. T., Nelson, S. F., Marques-Bonet, T., Ostrander, E. A., Wayne, R. K., and Novembre, J. (2016). Demographically-Based Evaluation of Genomic Regions under Selection in Domestic Dogs. PLoS Genetics 12(3), e1005851.
| Crossref | Google Scholar | PubMed |

Galibert, F., Quignon, P., Hitte, C., and André, C. (2011). Toward understanding dog evolutionary and domestication history. Comptes Rendus Biologies 334(3), 190-196.
| Crossref | Google Scholar | PubMed |

Gaunitz, C., Fages, A., Hanghøj, K., Albrechtsen, A., Khan, N., Schubert, M., Seguin-Orlando, A., Owens, I. J., Felkel, S., Bignon-Lau, O., De Barros Damgaard, P., Mittnik, A., Mohaseb, A. F., Davoudi, H., Alquraishi, S., Alfarhan, A. H., Al-Rasheid, K., Crubézy, E., Benecke, N., Olsen, S., Brown, D., Anthony, D., Massy, K., Pitulko, V., Kasparov, A., Brem, G., Hofreiter, M., Mukhtarova, G., Baimukhanov, N., Lõugas, L., Onar, V., Stockhammer, P. W., Krause, J., Boldgiv, B., Undrakhbold, S., Erdenebaatar, D., Lepetz, S., Mashkour, M., Ludwig, A., Wallner, B., Merz, V., Merz, I., Zaibert, V., Willerslev, E., Librado, P., Outram, A. K., and Orlando, L. (2018). Ancient genomes revisit the ancestry of domestic and Przewalski’s horses. Science 360(6384), 111-114.
| Crossref | Google Scholar | PubMed |

Gentry, A., Clutton-Brock, J., and Groves, C. P. (2004). The naming of wild animal species and their domestic derivatives. Journal of Archaeological Science 31(5), 645-651.
| Crossref | Google Scholar |

Gojobori, J., Arakawa, N., Xiaokaiti, X., Matsumoto, Y., Matsumura, S., Hongo, H., Ishiguro, N., and Terai, Y. (2024). Japanese wolves are most closely related to dogs and share DNA with East Eurasian dogs. Nature Communications 15(1), 1680.
| Crossref | Google Scholar | PubMed |

Goolmeer, T., Skroblin, A., Grant, C., van Leeuwen, S., Archer, R., Gore-Birch, C., and Wintle, B. A (2022). Recognizing culturally significant species and Indigenous-led management is key to meeting international biodiversity obligations. Conservation Letters 15, e12899.
| Crossref | Google Scholar |

Gopalakrishnan, S., Sinding, M.-H. S., Ramos-Madrigal, J., Niemann, J., Samaniego Castruita, J. A., Vieira, F. G., Carøe, C., Montero, M. d. M., Kuderna, L., Serres, A., González-Basallote, V. M., Liu, Y.-H., Wang, G.-D., Marques-Bonet, T., Mirarab, S., Fernandes, C., Gaubert, P., Koepfli, K.-P., Budd, J., Rueness, E. K., Sillero, C., Heide-Jørgensen, M. P., Petersen, B., Sicheritz-Ponten, T., Bachmann, L., Wiig, Ø., Hansen, A. J., and Gilbert, M. T. P. (2018). Interspecific Gene Flow Shaped the Evolution of the Genus Canis. Current Biology 28(21), 3441-3449.e5.
| Crossref | Google Scholar |

Greig, K., Gosling, A., Collins, C. J., Boocock, J., McDonald, K., Addison, D. J., Allen, M. S., David, B., Gibbs, M., Higham, C. F. W., Liu, F., McNiven, I. J., O’Connor, S., Tsang, C. H., Walter, R., and Matisoo-Smith, E. (2018). Complex history of dog (Canis familiaris) origins and translocations in the Pacific revealed by ancient mitogenomes. Scientific Reports 8(1), 9130.
| Crossref | Google Scholar | PubMed |

Heinsohn, T. (2014). Animal translocation:long-term human influences on the vertebrate zoogeography of Australasia (natural dispersal versus ethnophoresy). Australian Zoologist 32(3), 351-376.
| Crossref | Google Scholar |

Hofman, C. A., and Rick, T. C. (2018). Ancient Biological Invasions and Island Ecosystems: Tracking Translocations of Wild Plants and Animals. Journal of Archaeological Research 26(1), 65-115.
| Crossref | Google Scholar |

ICZN (1999). International Code of Zoological Nomenclature, 4th edn. (International Trust for Zoological Nomenclature: London, U.K.) Available at http://www.iczn.org/iczn/index

Jackson, S. M., Groves, C. P., Fleming, P. J. S., Aplin, K. P., Eldridge, M. D. B., Gonzalez, A., and Helgen, K. M. (2017). The wayward dog: is the Australian native dog or dingo a distinct species? Zootaxa 4317(2), 201-224.
| Crossref | Google Scholar |

Jackson, S. M., Fleming, P. J. S., Eldridge, M. D. B., Ingleby, S., Flannery, T., Johnson, R. N., Cooper, S. J. B., Mitchell, K. J., Souilmi, Y., Cooper, A., Wilson, D. E., and Helgen, K. M. (2019). The Dogma of Dingoes-Taxonomic status of the dingo: A reply to Smith et al. Zootaxa 4564(1), 198-212.
| Crossref | Google Scholar | PubMed |

Jackson, S. M., Fleming, P. J. S., Eldridge, M. D. B., Archer, M., Ingleby, S., Johnson, R. N., and Helgen, K. M. (2020). Taxonomy of the Dingo: It’s an ancient dog. Australian Zoologist 41, 347-357.
| Crossref | Google Scholar |

King, C., and Forsyth, D. (Eds.) (2021). ‘The Handbook of New Zealand Mammals.’ (CSIRO Publishing.)

Koungoulos, L. (2021). Domestication Through Dingo Eyes: An Australian Perspective on Human-Canid Interactions Leading to the Earliest Dogs. Human Ecology 49, 691-705.
| Crossref | Google Scholar |

Koungoulos, L., and Fillios, M. (2020). Hunting dogs down under? On the Aboriginal use of tame dingoes in dietary game acquisition and its relevance to Australian prehistory. Journal of Anthropological Archaeology 58, 101146.
| Google Scholar |

Koungoulos, L. G., Hulme-Beaman, A., Fillios, M., and Willandra Lakes Region World Heritage Aboriginal Advisory Group (2024). Phenotypic diversity in early Australian dingoes revealed by traditional and 3D geometric morphometric analysis. Scientific Reports 14(1),.
| Crossref | Google Scholar |

Kreplins, T. L., Stephens, D., Laing, N., and Oskam, C. (2024). An assessment of dingo ancestry in camp dogs in Western Australia. Australian Journal of Zoology 72(3), ZO24008.
| Crossref | Google Scholar |

Krofel, M., Hatlauf, J., Bogdanowicz, W., Campbell, L. A. D., Godinho, R., Jhala, Y. V., Kitchener, A. C., Koepfli, K. P., Moehlman, P., Senn, H., Sillero-Zubiri, C., Viranta, S., Werhahn, G., and Alvares, F. (2022). Towards resolving taxonomic uncertainties in wolf, dog and jackal lineages of Africa, Eurasia and Australasia. Journal of Zoology 316(3), 155-168.
| Crossref | Google Scholar |

Kumar, M., Conroy, G., Ogbourne, S., Cairns, K., Borburgh, L., and Subramanian, S. (2023). Genomic signatures of bottleneck and founder effects in dingoes. Ecology and Evolution 13(9), e10525.
| Crossref | Google Scholar | PubMed |

Larson, G., and Burger, J. (2013). A population genetics view of animal domestication. Trends in Genetics 29(4), 197-205.
| Crossref | Google Scholar | PubMed |

Letnic, M., Ritchie, E. G., and Dickman, C. R. (2012). Top predators as biodiversity regulators: the dingo Canis lupus dingo as a case study. Biological Reviews 87(2), 390-413.
| Crossref | Google Scholar | PubMed |

Lord, K., Feinstein, M., Smith, B., and Coppinger, R. (2013). Variation in reproductive traits of members of the genus Canis with special attention to the domestic dog (Canis familiaris). Behavioural Processes 92(0), 131-142.
| Crossref | Google Scholar | PubMed |

Lord, K. A., Larson, G., Allaby, R. G., and Karlsson, E. K. (2025). A universally applicable definition for domestication. Proceedings of the National Academy of Sciences of the United States of America 122(22), e2413207122.
| Crossref | Google Scholar | PubMed |

Manwell, C., and Baker, C. M. A. (1984). Domestication of the dog: hunter, food, bed-warmer, or emotional object? Zeitschrift für Tierzüchtung und Züchtungsbiologie 101(1–5), 241-256.
| Crossref | Google Scholar |

Marshall, F. B., Dobney, K., Denham, T., and Capriles, J. M. (2014). Evaluating the roles of directed breeding and gene flow in animal domestication. Proceedings of the National Academy of Sciences of the United States of America 111(17), 6153-6158.
| Crossref | Google Scholar | PubMed |

Martins, R. F., Schmidt, A., Lenz, D., Wilting, A., and Fickel, J. (2018). Human‐mediated introduction of introgressed deer across Wallace’s line: Historical biogeography of Rusa unicolor and R. timorensis. Ecology and Evolution 8(3), 1465-1479.
| Crossref | Google Scholar | PubMed |

National Academies of Sciences, Engineering, and Medicine (2019). ‘Evaluating the Taxonomic Status of the Mexican Gray Wolf and the Red Wolf.’ (The National Academies Press: Washington, DC.) 10.17226/25351

Oetjens, M. T., Martin, A., Veeramah, K. R., and Kidd, J. M. (2018). Analysis of the canid Y-chromosome phylogeny using short-read sequencing data reveals the presence of distinct haplogroups among Neolithic European dogs. BMC Genomics 19(1), 350.
| Crossref | Google Scholar | PubMed |

Oskarsson, M. C. R., Klütsch, C. F. C., Boonyaprakob, U., Wilton, A., Tanabe, Y., and Savolainen, P. (2012). Mitochondrial DNA data indicate an introduction through Mainland Southeast Asia for Australian dingoes and Polynesian domestic dogs. Proceedings of the Royal Society B: Biological Sciences 279(1730), 967-974.
| Crossref | Google Scholar | PubMed |

Parker, H. G., Dreger, D. L., Rimbault, M., Davis, B. W., Mullen, A. B., Carpintero-Ramirez, G., and Ostrander, E. A. (2017). Genomic Analyses Reveal the Influence of Geographic Origin, Migration, and Hybridization on Modern Dog Breed Development. Cell Reports 19(4), 697-708.
| Crossref | Google Scholar | PubMed |

Pendleton, A. L., Shen, F., Taravella, A. M., Emery, S., Veeramah, K. R., Boyko, A. R., and Kidd, J. M. (2018). Comparison of village dog and wolf genomes highlights the role of the neural crest in dog domestication. BMC Biology 16(1), 64.
| Crossref | Google Scholar | PubMed |

Perri, A. R., Feuerborn, T. R., Frantz, L. A. F., Larson, G., Malhi, R. S., Meltzer, D. J., and Witt, K. E. (2021). Dog domestication and the dual dispersal of people and dogs into the Americas. Proceedings of the National Academy of Sciences of the United States of America 118(6), e2010083118.
| Crossref | Google Scholar | PubMed |

Pilot, M. (2021). Disentangling the admixed trails of grey wolf evolution. Molecular Ecology 30(24), 6509-6512.
| Crossref | Google Scholar | PubMed |

Pilot, M., Malewski, T., Moura, A. E., Grzybowski, T., Oleński, K., Ruść, A., Kamiński, S., Fadel, F. R., Mills, D. S., Alagaili, A. N., Mohammed, O. B., Kłys, G., Okhlopkov, I. M., Suchecka, E., and Bogdanowicz, W. (2015). On the origin of mongrels: evolutionary history of free-breeding dogs in Eurasia. Proceedings of the Royal Society B: Biological Sciences 282(1820), 20152189.
| Crossref | Google Scholar | PubMed |

Purugganan, M. D. (2022). What is domestication? Trends in Ecology & Evolution 37(8), 663-671.
| Crossref | Google Scholar | PubMed |

Ritchie, E. G., Smith, B. P., van Eeden, L. M., and Nimmo, D. G. (2018). Species definitions shape policy. Science 361(6409), 1324.
| Crossref | Google Scholar | PubMed |

Sacks, B. N., Brown, S. K., Stephens, D., Pedersen, N. C., Wu, J.-T., and Berry, O. (2013). Y chromosome analysis of dingoes and Southeast Asian village dogs suggests a Neolithic continental expansion from Southeast Asia followed by multiple Austronesian dispersals. Molecular Biology and Evolution 30, 1103-18.
| Crossref | Google Scholar | PubMed |

Savolainen, P., Leitner, T., Wilton, A. N., Matisoo-Smith, E., and Lundeberg, J. (2004). A detailed picture of the origin of the Australian dingo, obtained from the study of mitochondrial DNA. Proceedings of the National Academy of Sciences 101(33), 12387-12390.
| Crossref | Google Scholar |

Scarsbrook, L., Cairns, K. M., Mitchell, K. J., Bougiouri, K., Evin, A., Harris, A. C., Wood, A. E., Zehui, Zhang, Z., Alves, J. M., Lawson, D. J., Ditchfield, P. W., Granja-Martins, S., Styring, A. K., Tabadda, K., Thalmann, O., Crowther, M. S., Curry, M., Feuerborn, T. R., Koungoulos, L. G., Letnic, M., Reed, E. H., Sabin, R., Parker, H. G., Ostrander, E. A., Frantz, L. A. F., Larson, G., and Fillios, M. A. (In Press). The Impacts of European Arrival on Australian Dingoes. Proceedings of the National Academy of Sciences. Accepted 09/07/2025.

Schweizer, R. M., Durvasula, A., Smith, J., Vohr, S. H., Stahler, D. R., Galaverni, M., Thalmann, O., Smith, D. W., Randi, E., Ostrander, E. A., Green, R. E., Lohmueller, K. E., Novembre, J., and Wayne, R. K. (2018). Natural Selection and Origin of a Melanistic Allele in North American Gray Wolves. Molecular Biology and Evolution 35(5), 1190-1209.
| Crossref | Google Scholar |

Shannon, L. M., Boyko, R. H., Castelhano, M., Corey, E., Hayward, J. J., McLean, C., White, M. E., Abi Said, M., Anita, B. A., Bondjengo, N. I., Calero, J., Galov, A., Hedimbi, M., Imam, B., Khalap, R., Lally, D., Masta, A., Oliveira, K. C., Pérez, L., Randall, J., Tam, N. M., Trujillo-Cornejo, F. J., Valeriano, C., Sutter, N. B., Todhunter, R. J., Bustamante, C. D., and Boyko, A. R. (2015). Genetic structure in village dogs reveals a Central Asian domestication origin. Proceedings of the National Academy of Sciences of the United States of America 112(44), 13639-13644.
| Crossref | Google Scholar | PubMed |

Shipman, P. (2021). What the dingo says about dog domestication. The Anatomical Record 304(1), 19-30.
| Crossref | Google Scholar | PubMed |

Sillero-Zubiri, C. (2009. Family Canidae (Dogs). In ‘Handbook of the Mammals of the World,Vol. 1. Carnivores’. Lynx Edn. (Eds D. E. Wilson, and R. A. Mittermeier.) pp. 352–466. (Barcelon.)

Sinding, M.-H. S., Gopalakrishnan, S., Ramos-Madrigal, J., de Manuel, M., Pitulko, V. V., Kuderna, L., Feuerborn, T. R., Frantz, L. A. F., Vieira, F. G., Niemann, J., Samaniego Castruita, J. A., Carøe, C., Andersen-Ranberg, E. U., Jordan, P. D., Pavlova, E. Y., Nikolskiy, P. A., Kasparov, A. K., Ivanova, V. V., Willerslev, E., Skoglund, P., Fredholm, M., Wennerberg, S. E., Heide-Jørgensen, M. P., Dietz, R., Sonne, C., Meldgaard, M., Dalén, L., Larson, G., Petersen, B., Sicheritz-Pontén, T., Bachmann, L., Wiig, Ø., Marques-Bonet, T., Hansen, A. J., and Gilbert, M. T. P. (2020). Arctic-adapted dogs emerged at the Pleistocene–Holocene transition. Science 368(6498), 1495-1499.
| Crossref | Google Scholar | PubMed |

Skoglund, P., Ersmark, E., Palkopoulou, E., and Dalén, L. (2015). Ancient wolf genome reveals an early divergence of domestic dog ancestors and admixture into high-latitude breeds. Current Biology 25(11), 1515-1519.
| Crossref | Google Scholar | PubMed |

Smith, B. P., and Litchfield, C. A. (2009). A Review of the Relationship between Indigenous Australians, Dingoes (Canis dingo) and Domestic Dogs (Canis familiaris). Anthrozoös 22(2), 111-128.
| Google Scholar |

Smith, B. P., Cairns, K. M., Adams, J. W., Newsome, T. M., Fillios, M., Déaux, E. C., Parr, W. C. H., Letnic, M., van Eeden, L. M., Appleby, R. G., Bradshaw, C., Savolainen, P., Ritchie, E. G., Nimmo, D. G., Archer-Lean, C., Greenville, A. C., Dickman, C. R., Watson, L., Moseby, K. E., Doherty, T. S., Wallach, A. D., Morrant, D. S., and Crowther, M. S. (2019). Taxonomic status of the Australian dingo: the case for Canis dingo Meyer, 1793. Zootaxa 4564(1), 173-197.
| Crossref | Google Scholar | PubMed |

Smith, B. P., Appleby, R. G., and Cairns, K. M. (2023). Reimagining the Dingo: The ‘Australian Wolf ’ or Just a Feral Dog?. In ‘The Wolf’. (Eds I. Convery, O. Nevin, E. van Maanen, et al) pp. 161-178. (Boydell and Brewer.) 10.1515/9781805430551-020

Souilmi, Y., Wasef, S., Williams, M. P., Conroy, G., Bar, I., Bover, P., Dann, J., Heiniger, H., Llamas, B., Ogbourne, S., Archer, M., Ballard, J. W. O., Reed, E., Tobler, R., Koungoulos, L., Walshe, K., Wright, J. L., Balme, J., O’Connor, S., Cooper, A., and Mitchell, K. J. (2024). Ancient genomes reveal over two thousand years of dingo population structure. Proceedings of the National Academy of Sciences of the United States of America 121(30), e2407584121.
| Crossref | Google Scholar | PubMed |

Stephens, D., Wilton, A. N., Fleming, P. J. S., and Berry, O. (2015). Death by sex in an Australian icon: a continent-wide survey reveals extensive hybridization between dingoes and domestic dogs. Molecular Ecology 24(22), 5643-5656.
| Crossref | Google Scholar | PubMed |

Stephens, D., Fleming, P. J. S., Sawyers, E., and Mayr, T. P. (2022). An isolated population reveals greater genetic structuring of the Australian dingo. Scientific Reports 12(1), 19105.
| Crossref | Google Scholar | PubMed |

Surbakti, S., Parker, H. G., McIntyre, J. K., Maury, H. K., Cairns, K. M., Selvig, M., Pangau-Adam, M., Safonpo, A., Numberi, L., Runtuboi, D. Y. P., Davis, B. W., and Ostrander, E. A. (2020). New Guinea highland wild dogs are the original New Guinea singing dogs. Proceedings of the National Academy of Sciences of the United States of America 117(39), 24369-24376.
| Crossref | Google Scholar | PubMed |

Thalmann, O., Shapiro, B., Cui, P., Schuenemann, V. J., Sawyer, S. K., Greenfield, D. L., Germonpré, M. B., Sablin, M. V., López-Giráldez, F., Domingo-Roura, X., Napierala, H., Uerpmann, H.-P., Loponte, D. M., Acosta, A. A., Giemsch, L., Schmitz, R. W., Worthington, B., Buikstra, J. E., Druzhkova, A., Graphodatsky, A. S., Ovodov, N. D., Wahlberg, N., Freedman, A. H., Schweizer, R. M., Koepfli, K.-P., Leonard, J. A., Meyer, M., Krause, J., Pääbo, S., Green, R. E., and Wayne, R. K. (2013). Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs. Science 342(6160), 871-874.
| Crossref | Google Scholar | PubMed |

Thiele, K. R., Conix, S., Pyle, R. L., Barik, S. K., Christidis, L., Costello, M. J., van Dijk, P. P., Kirk, P., Lien, A., Thomson, S. A., Zachos, F. E., Zhang, Z.-Q., and Garnett, S. T. (2021). Towards a global list of accepted species I. Why taxonomists sometimes disagree, and why this matters. Organisms Diversity & Evolution 21(4), 615-622.
| Crossref | Google Scholar |

Thomson, P. C., Rose, K., and Kok, N. E. (1992). The behavioural ecology of dingoes in north-Western Australia. V. Population dynamics and variation in the social system. Wildlife Research 19(5), 565-583.
| Google Scholar |

Veron, G., Willsch, M., Dacosta, V., Patou, M.-L., Seymour, A., Bonillo, C., Couloux, A., Wong, S. T., Jennings, A. P., Fickel, J., and Wilting, A. (2014). The distribution of the Malay civet Viverra tangalunga (Carnivora: Viverridae) across Southeast Asia: natural or human-mediated dispersal? Zoological Journal of the Linnean Society 170(4), 917-932.
| Crossref | Google Scholar |

vonHoldt, B. M., and Aardema, M. L. (2020). Updating the bibliography of interbreeding among Canis in North America. Journal of Heredity 111, 249-262.
| Crossref | Google Scholar | PubMed |

vonHoldt, B. M., Pollinger, J. P., Lohmueller, K. E., Han, E., Parker, H. G., Quignon, P., Degenhardt, J. D., Boyko, A. R., Earl, D. A., Auton, A., Reynolds, A., Bryc, K., Brisbin, A., Knowles, J. C., Mosher, D. S., Spady, T. C., Elkahloun, A., Geffen, E., Pilot, M., Jedrzejewski, W., Greco, C., Randi, E., Bannasch, D., Wilton, A., Shearman, J., Musiani, M., Cargill, M., Jones, P. G., Qian, Z., Huang, W., Ding, Z.-L., Zhang, Y.-p., Bustamante, C. D., Ostrander, E. A., Novembre, J., and Wayne, R. K. (2010). Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication. Nature 464, 898-902.
| Crossref | Google Scholar | PubMed |

vonHoldt, B., Pollinger, J., Earl, D., Parker, H., Ostrander, E., and Wayne, R. (2013). Identification of recent hybridization between gray wolves and domesticated dogs by SNP genotyping. Mammalian Genome 24(1–2), 80-88.
| Crossref | Google Scholar | PubMed |

vonHoldt, B. M., Brzeski, K. E., Wilcove, D. S., and Rutledge, L. Y. (2018). Redefining the role of admixture and genomics in species conservation. Conservation Letters 11(2), e12371.
| Crossref | Google Scholar |

Weeks, A. R., Kriesner, P., Bartonicek, N., van Rooyen, A., Cairns, K. M., and Ahrens, C. W. (2024). Genetic structure and common ancestry expose the dingo-dog hybrid myth. Evolution Letters 9, 1-12.
| Crossref | Google Scholar | PubMed |

Zachos, F. E. (2016) ‘Species Concepts in Biology: Historical Development, Theoretical Foundations and Practical Relevance.’ (Springer: Charm, Switzerland.)

Zeder, M. A. (2012). The Domestication of Animals. Journal of Anthropological Research 68(2), 161-190.
| Crossref | Google Scholar |

Zeller, U., and Göttert, T. (2019). The relations between evolution and domestication reconsidered - Implications for systematics, ecology, and nature conservation. Global Ecology and Conservation 20, e00756.
| Crossref | Google Scholar |

Zhang, S.-j., Wang, G.-D., Ma, P., Zhang, L.-l., Yin, T.-T., Liu, Y.-h., Otecko, N. O., Wang, M., Ma, Y.-p., Wang, L., Mao, B., Savolainen, P., and Zhang, Y.-p. (2020). Genomic regions under selection in the feralization of the dingoes. Nature Communications 11(1), 671.
| Crossref | Google Scholar | PubMed |