The myth of wild dogs in Australia: are there any out there?

Abstract

Hybridisation between wild and domestic canids is a global conservation and management issue. In Australia, dingoes are a distinct lineage of wild-living canid with a controversial domestication status. They are mainland Australia’s apex terrestrial predator. There is ongoing concern that the identity of dingoes has been threatened from breeding with domestic dogs, and that feral dogs have established populations in rural Australia. We collate the results of microsatellite DNA testing from 5039 wild canids to explore patterns of domestic dog ancestry in dingoes and observations of feral domestic dogs across the continent. Only 31 feral dogs were detected, challenging the perception that feral dogs are widespread in Australia. First generation dingo × dog hybrids were similarly rare, with only 27 individuals identified. Spatial patterns of genetic ancestry across Australia identified that dingo populations in northern, western and central Australia were largely free from domestic dog introgression. Our findings challenge the perception that dingoes are virtually extinct in the wild and that feral dogs are common. A shift in terminology from wild dog to dingo would better reflect the identity of these wild canids and allow more nuanced debate about the balance between conservation and management of dingoes in Australia.

Full text

The myth of wild dogs in Australia: are there any out there?
Kylie M. Cairns
A
,
B
,
E
,Mathew S. Crowther
C
,Bradley Nesbitt
D
and
Mike Letnic
A
,
B
A
Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences,
University of New South Wales, Sydney, NSW 2052, Australia.
B
Evolution and Ecology Research Centre, School of Biological, Earth and Environmental Sciences,
University of New South Wales, Sydney, NSW 2052, Australia.
C
School of Life and Environmental Sciences, University of Sydney, NSW 2006, Australia.
D
School of Environmental and Rural Science, University of New England, Armidale, NSW 2350, Australia.
E
Corresponding author. Email: k.cairns@unsw.edu.au
Abstract. Hybridisation between wild and domestic canids is a global conservation and management issue. In Australia,
dingoes are a distinct lineage of wild-living canid with a controversial domestication status. They are mainland Australia’s
apex terrestrial predator. There is ongoing concern that the identity of dingoes has been threatened from breeding with
domestic dogs, and that feral dogs have established populations in rural Australia. We collate the results of microsatellite
DNA testing from 5039 wild canids to explore patterns of domestic dog ancestry in dingoes and observations of feral
domestic dogs across the continent. Only 31 feral dogs were detected, challenging the perception that feral dogs are
widespread in Australia. First generation dingo dog hybrids were similarly rare, with only 27 individuals identified.
Spatial patterns of genetic ancestry across Australia identified that dingo populations in northern, western and central
Australia were largely free from domestic dog introgression. Our findings challenge the perception that dingoes are
virtually extinct in the wild and that feral dogs are common. A shift in terminology from wild dog to dingo would better
reflect the identity of these wild canids and allow more nuanced debate about the balance between conservation and
management of dingoes in Australia.
Keywords: admixture, Australia, Canis dingo,Canis familiaris, dingo, dog, domestication, feral dog, introgression, wild dog.
Received 17 August 2020, accepted 26 February 2021, published online 26 March 2021
Introduction
The occurrence of feral domestic dogs is rare, and distinct from
the close to a billion free-breeding or village dogs that exist
globally (Gompper 2013;Pilot et al. 2015). Free-breeding or
village dogs are those that live in and around human settlements,
rely upon anthropogenic food or water sources, breed freely with
each other, and are not owned or cared for by people (Hughes
and Macdonald 2013). Feral dogs are those that are living in a
wild state not in the vicinity of human settlements: they may be
escaped pets or self-sustaining populations. Empirical data from
remote living free-breeding dog populations suggests these
populations rely upon recruitment from stray or owned dogs
because their reproductive success is low, i.e. pups rarely
survive past 1 year (Boitani et al. 1995,2006). The only
acknowledged example of a true wild-living self-sustaining feral
dog population occurred in the Gala´pagos; it was founded by a
variety of breeds in the 1800s and persisted until the 1980s
(Barnett 1986;Reponen et al. 2014). Despite there being a
robust population of free-breeding or stray dogs associated with
towns in the Gala´pagos, there was limited evidence of mixing
between the free-breeding dog and feral dog populations based
on genetic analysis (Reponen et al. 2014). The Gala´pagos feral
dog population was eradicated using 1080 poisoning of water
and meat baits in the 1980s (Barnett 1986) and the population
did not re-establish despite the presence of free-breeding and pet
dogs in nearby human settlements.
Dingoes, including New Guinea singing dogs, form a discrete
lineage from Eurasian and modern domestic dogs (Bergstro¨m
et al. 2020;Surbakti et al. 2020;Cairns 2021). Their domestica-
tion status and taxonomic nomenclature is disputed, with some
considering them Canis familiaris, a feral domestic dog (Jackson
et al. 2017,2019) and others calling them Canis dingo, a wild
protodog (Crowther et al. 2014;Smith et al. 2019;Zhang et al.
2020). Globally, most free-breeding, village and breed dogs fall
within the modern domestic dog lineage (Bergstro¨m et al. 2020;
Surbakti et al. 2020;Cairns 2021). A close relationship between
dingoes, Asian wolves, and some East Asian dogs has been
observed, suggesting the dingoes’ ancestor was of Asian origin
(vonHoldt et al. 2010;Oskarsson et al. 2011;Freedman et al.
2014;Surbakti et al. 2020;Zhang et al. 2020;Cairns 2021).
Dingoes and New Guineasinging dogs are examples of true wild-
living dogs that are not reliant on artificial water or food sources.
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Dingoes fill the ecological role of terrestrial apex predator on
mainland Australia (Newsome et al. 2001;Letnic and Koch
2010;Letnic et al. 2012;Letnic et al. 2013;Morrant et al. 2017).
Molecular dating indicates that dingoes and New Guinea singing
dogs diverged from their ancestral population approximately
8000–12 000 years ago (Cairns and Wilton 2016;Cairns et al.
2017;Zhang et al. 2020). Dingoes remained reproductively
isolated from domestic dogs until 1788.
There is rising global concern about the occurrence of
hybridisation between wild and domestic canids or sympatric
wild canids (Gopalakrishnan et al. 2018;Salvatori et al. 2020;
vonHoldt and Aardema 2020). Hybridisation is the process of
interbreeding between two species or varieties, generally F1
offspring would be referred to as hybrids and the offspring of F1
hybrids with animals from a parental species or variety would be
called backcrosses. Interbreeding between varieties or species
results in the exchange or mixing of genetic material (genetic
admixture). The transfer of genetic material from one species
into another through hybridisation and backcrossing is called
introgression (Harrison and Larson 2014). The occurrence of
genetic admixture may be modern or historical, and in some
cases is the result of anthropogenic actions. In canids, the
phenomenon of interspecific introgression has been observed
between species such as grey wolves and dogs (Vila` and Wayne
1999;Anderson et al. 2009;vonHoldt et al. 2011,2016;
Schweizer et al. 2018), coyotes and wolves (Bohling et al.
2016;vonHoldt et al. 2016), red wolves and coyotes (Miller
et al. 2003;Adams et al. 2007;Schmutz et al. 2007;Bohling and
Waits 2015), jackals and dogs (Galov et al. 2015), dingoes and
dogs (Newsome and Corbett 1985;Wilton 2001;Claridge et al.
2014;Stephens et al. 2015;Cairns et al. 2019).
One of the concerns raised by hybridisation is genetic swamp-
ing, whereby the genetic identity of a population is threatened by
introgression of genes from another population or species. For
example, the Scottish wildcat is threatened by hybridisation and
subsequent introgression from domestic cats to the extent that
contemporary wildcat populations exhibit extensive levels of
domestic cat ancestry (Daniels et al. 1998;Kitchener et al. 2005;
Macdonald et al. 2010). Indeed, most wildcats in Scotland carry
significant domestic cat ancestry, and the occurrence of hybridisa-
tion is believed to have accelerated in the last 50–100 years
(Mattucci et al. 2019;Senn et al. 2019). Similar concerns have
been raised in Australia with many dingoes, particularly in
southeastern Australia, exhibiting genetic, morphological or
phenotypic evidence of domestic dog ancestry (Newsome and
Corbett 1985;Daniels and Corbett 2003;Jones 2009;Stephens
et al. 2015). There is also widespread concern that feral domestic
dogs have established in the wild across Australia (Fleming et al.
2001;NSW Threatened Species Scientific Committee 2009).
Since the 1980s rising concern about domestic dog ancestry
and the occurrence of hybridisation events has led to shifts in the
management and conservation status of dingoes but also a
duality in their identity. For example, in Victoria dingoes are
now listed as a threatened species, but wild dogs are listed as a
declared pest, where wild dogs are defined as feral or wild
populations of dogs (Canis familiaris) and dingo dog hybrids
(Canis dingo Canis familiaris)(DEPI 2013). In New South
Wales (NSW), the listing of ‘predation and hybridisation by
feral dogs (Canis familiaris)’ as a key threatening process
implies that dingoes are ‘under serious decline as a consequence
of hybridisation’ (NSW Threatened Species Scientific Commit-
tee 2009). Indeed, there has been concern in NSW that feral dogs
and dingo dog hybrids with low levels of dingo ancestry have
essentially replaced dingoes in the wild (Claridge et al. 2014;
Stephens et al. 2015). For example, the NSW key threatening
process determination states that ‘due to the constant influx of
Domestic Dogs into natural ecosystems, lasting eradication of
even local populations of Feral Dogs is difficult’ (NSW Threat-
ened Species Scientific Committee 2009). Accordingly, the term
‘wild dog’ is now used ubiquitously by state government and pest
control organisations when communicating about management
programs directed at controlling wild canids (Letnic 2012;
Kreplins et al. 2019). However, it is clear from social studies of
public perception and also expectations about the management of
dingoes vs wild dogs, that the general public believe the term wild
dog refers only to feral dogs and does not properly understand that
the term wild dog is defined as including dingoes, dingo dog
hybrids and feral dogs (van Eeden et al. 2020).
Before the 2000s a majority of our knowledge about
dingo dog hybridisation was based on assessment of skull
morphology and physical appearance (Newsome et al. 1980;
Newsome and Corbett 1985;Jones 1990;Corbett 2001;Fleming
et al. 2001). A microsatellite DNA test for assessing the ancestry
of dingoes was developed in 1999 (Wilton et al. 1999;Wilton
2001) and has since become widely used by wildlife managers
and conservation groups. Studies assessing the reliability of
morphological, physical and genetic methods of ancestry assess-
ment in dingoes have highlighted the importance of using
genetic data (Elledge et al. 2008;Parr et al. 2016). Stephens
et al. (2015) undertook microsatellite DNA testing of wild
canids across Australia and identified regional patterns of
domestic dog introgression in dingoes, with dog ancestry par-
ticularly prevalent in southeastern Australia. However, a major
limitation of Stephens et al. (2015) is the low number of samples
(95) from NSW. A more detailed study using the same genetic
markers and significantly higher density of sampling across
northeastern NSW identified several key hotspots of high dingo
ancestry (Cairns et al. 2019). Their finding that a majority of
wild dingoes in NSW were pure dingoes or carried more than
75% dingo ancestry is a stark contrast to the common public
perception that feral dogs are widespread and established in the
wild (NSW Threatened Species Scientific Committee 2009;
Claridge et al. 2014;ABC Landline 2019). Here we collate
and analyse genetic ancestry data based on microsatellite DNA
testing from 5039 samples to examine the occurrence of feral
dogs and F1 or F2 dingo dog hybrids across Australia.
Critically, this study includes a broader set of samples from
southeastern Australia including from southern NSW filling
knowledge gaps about the ancestry and identity of wild canids
in southeastern Australia. Spatial patterns of domestic dog
introgression across Australia are also examined using the
dataset. These data inform ongoing debate about the appropriate
terminology and management of wild canids in Australia.
Methods and materials
DNA testing based on a widely used 23 microsatellite marker set
was used to estimate dingo ancestry in Stephens et al. (2015),
BAustralian Mammalogy K. M. Cairns et al.

Cairns et al. (2019) and a previously unpublished dataset (available
in the BioStudies database under accession number S-BSST501
https://www.ebi.ac.uk/biostudies/studies/S-BSST501). Briefly, a
panel of 23 microsatellites were amplified and genotyped in
the wild canid samples based on the original methodology of
Wilton (2001) and Elledge (2008). Ancestry modelling was
performed in STRUCTURE with reference populations of
known dingoes and dogs using the admixture and correlated
allele frequency models. Cairns et al. (2019) and the unpub-
lished dataset used a set of 50 dingoes and 66 mixed breed dogs
as reference populations; to account for regional variation these
analyses included a set of 13 wild dingoes from northern and
western Australia. Stephens et al. (2015) used a set of 322 post-
priori identified reference dingoes and 102 domestic dogs. In all
three studies simulations were run with 200 000 iterations, a
20 000 iteration burn-in period and 10 replicates of each K ¼2
was performed. Previously, modelling demonstrated that K ¼2
was the appropriate model for assessing ancestry in Australian
wild canids and modelling was run with the USEPOPINFO flag
to allow population allele frequencies to be updated only from
the defined reference population individuals (Stephens et al.
2015;Cairns et al. 2019). STRUCTURE reports estimated
ancestry proportions (q-values) for each genetic cluster for each
sample (Stephens et al. 2015;Cairns et al. 2019). In a K ¼2
analysis each individual has a q-value for the domestic dog
cluster and for the dingo cluster. The dingo cluster q-value is
used to define animals as either a pure dingo, probable dingo,
dingo with .75% ancestry, dingo with 65–75% ancestry, dingo
with 50–65% ancestry, feral dog hybrid or feral domestic dog
(Table 1;Stephens et al. 2015;Cairns et al. 2019). We define
feral dog hybrids with a q-value between 0.25 and 0.49 to be
possible F1 or F2 dingo dog hybrids (Stephens et al. 2015;
Cairns et al. 2019). An F1 hybrid is defined as the offspring of a
dingo a dog and an F2 hybrid is the offspring of two F1
dingo dog hybrids.
Raw microsatellite data could not be compared because of
slightly different microsatellite amplification conditions. To
confirm that ancestry estimates were equivalent between Cairns
et al. (2019) and Stephens et al. (2015), a set of 13 wild canids
were genotyped and ancestry estimates calculated by both
laboratories (Table 2).
Between the three-studies DNA ancestry estimates from 5039
wild canids collected by trappers, wildlife managers and govern-
ment agencies across public and private lands in Australia were
reported. A majority of the wild canids were trapped/shot as part
of broadscale wild canid management to protect livestock from
predation. Samples from the three datasets were collected
between 1996 and 2014. We collated these DNA ancestry
estimates (STRUCTURE q-values) together with location coor-
dinates (Supplementary Material S1 dataset).
Table 1. Dingo purity categories and cut offs for average 3Q and STRUCTURE methods
Score Category Average 3Q
A
value and
doglike allele cut offs
STRUCTURE
B
average
q-value cut offs
1 Dingo 1 (dingo with no dog ancestry) 3Q .0.1 and no doglike alleles 1.0–0.90
2 Dingo 2 (likely dingo with no dog ancestry) 0.05 ,3Q ,0.1 and #1 doglike alleles 0.89–0.80
3 Dingo with dog ancestry 1 (.75% dingo) 0 ,3Q ,0.05 0.79–0.70
4 Dingo with dog ancestry 2 (65–75% dingo) 0.1 ,3Q ,0 0.69–0.60
5 Dingo with dog ancestry 3 (50–64% dingo) 0.25 ,3Q ,0.1 0.59–0.50
6 Feral dog with dingo ancestry (,50% dingo) 0.5 ,3Q ,0.25 0.49–0.25
7 Feral dog 3Q ,0.5 0.24–0.0
A
According to Wilton (2001) and Wilton et al. (1999).
B
According to Stephens et al. (2015).
Table 2. Comparison of STRUCTURE q-value ancestry estimates for 13 repeated samples from Stephens et al. (2015) and Cairns et al. (2019)
Ancestry modelling from Cairns et al. (2019) Ancestry modelling from Stephens et al. (2015) Difference between
Cairns et al. (2019) and
Stephens et al. (2015) q-value
ID # markers dingo ancestry
(q-value)
ID # markers dingo ancestry
(q-value)
W0021 23 0.87 wdi0053 23 0.83 0.04
W0022 22 0.88 wdi0163 23 0.89 –0.01
W0023 22 0.94 wdi0627 23 0.93 0.01
W0024 23 0.92 wdi0840 21 0.99 –0.07
W0025 22 0.89 wdi1736 23 0.96 –0.07
W0026 22 0.92 wdi1811 23 0.89 0.03
W0027 23 0.92 wdi1884 23 0.94 –0.02
W0028 22 0.77 wdi2397 23 0.77 0.00
W0029 22 0.96 di0910 23 0.92 0.04
W0030 23 0.89 di1050 23 0.9 –0.01
W0031 23 0.89 di1270 23 0.88 0.01
W0033 22 0.94 de013 22 0.94 0.00
W0034 23 0.92 de017 22 0.94 –0.02
The wild dog myth Australian Mammalogy C

The distribution of feral dogs with no dingo ancestry and
possible F1 or F2 dingo dog hybrids was mapped using QGIS
ver. 3.01 (QGIS 2020). We also explore the distribution of
dingoes with varying degrees of dog ancestry across Australia as
follows: the location of 5039 samples with DNA ancestry results
were mapped in QGIS, a 0.3 0.3 degree hexagonal grid was
drawn and the mean and median ‘q-score’ of the samples within
each grid cell was calculated (using the join attributes by
location tool). We also mapped the occurrence of feral dogs
with no dingo ancestry and possible F1 or F2 dingo dog
hybrids across Australia.
Results
We collated the ancestry results of 3641 samples from Stephens
et al. (2015), 753 samples from Cairns et al. (2019) and 611
samples from the unpublished dataset. Ancestry estimates were
consistent between Stephens et al. (2015) and Cairns et al. (2019)
based on comparison of results for 13 repeated samples (Table 2).
Out of 5039 samples that were DNA tested the breakdown
of dingo ancestry results are as follows: 33.7% pure dingoes,
30.4% probable dingoes, 19.8% greater than75% dingo ancestry,
11.7% greater than 65% dingo ancestry and 3.2% greater than
50% dingo ancestry (Table 3). Feral dogs and F1 or F2 dingo-dog
hybrids were rarely collected from the wild and made up less
than 1.2% of the wild canid population (Table 3). In total, only
31 feral dogs with no evidence of dingo ancestry were observed
and27probableF1orF2dingodog hybrids (q-value between
0.25 and 0.49) were identified in the sample. The occurrence of
dog introgression differed between states (Fig. 1) and was more
prevalent in southeastern Australia.
Table 3. Ancestry of 5039 wild canids across Australia according to STRUCTURE modelling by Stephens et al. (2015),Cairns et al. (2019) and
this study
Category WA SA NT Qld NSW Vic. ACT Australia
Dingo 1 (dingo with
no dog ancestry)
61.9% (1414) 34.5% (51) 88.2% (112) 21.9% (78) 2.2% (29) 1.1% (8) 5.8% (6) 33.7% (1698)
Dingo 2 (likely dingo
with no dog ancestry)
35.0% (799) 56.8% (84) 10.2% (13) 46.6% (166) 23.8% (314) 16.8% (118) 35.6% (37) 30.4% (1531)
Dingo 1 and 2 (dingoes
with no dog ancestry
detected)
96.9% (2213) 91.2% (135) 98.4% (125) 68.5% (244) 26% (343) 17.9% (126) 41.3% (43) 64.1% (3229)
Dingo with dog ancestry
1(.75% dingo)
2.3% (52) 8.8% (13) 0.8% (1) 23.6% (84) 39.5% (521) 40.5% (284) 41.3% (43) 19.8% (998)
Dingo with dog ancestry
2 (65–75% dingo)
0.02% (1) 0 0.8% (1) 4.8% (17) 25.6% (337) 32.1% (225) 10.6% (11) 11.7% (592)
Dingo with dog ancestry
3 (50–64% dingo)
0.02% (1) 0 0 2.2% (8) 7.4% (97) 7.7% (54) 1.9% (2) 3.2% (162)
Feral dog with dingo
ancestry (,50% dingo)
0.2% (5) 0 0 0.6% (2) 0.9% (12) 1.1% (8) 0 0.5% (27)
Feral dog 0.5% (12) 0 0 0.3% (1) 0.6% (8) 0.7% (5) 4.8% (5) 0.6% (31)
WA
0
10
20
30
40
50
60
70
80
90
100
SA NT Qld NSW Vic. ACT
Feral dog (no dingo)
Feral dog (<50% dingo)
Dingo (>50% dingo)
Dingo (>65% dingo)
Dingo (>75% dingo)
Pure dingo (1 and 2)
Fig. 1. Proportion of Australian wild canids that were pure dingoes, dingoes with domestic dog introgression, possible F1 dingo dog
hybrids, and feral domestic dogs, by state.
DAustralian Mammalogy K. M. Cairns et al.

Mapping of wild canid ancestry across Australia indicates that
domestic dog introgression is largely restricted to southeastern
Australia (Fig. 2; Supplementary Material S2). Across northern,
central and western Australia,the dingo population is genetically
intact, i.e. with limited or no domestic dog introgression. In
southeastern Australia there are regions with dingo populations
that are genetically intact (Fig. 2) and most populations maintain
a dingo-dominant identity (median ancestry is greater than 75%
dingo). Feral dogs were restricted mostly to southeastern Aus-
tralia and were captured in relatively close proximity to human
settlements (Fig. 2). Interestingly, no feral dogs were identified in
central Australia.
Discussion
Extensive DNA testing across Australia detected very few feral
dogs. Out of the 5039 wild canids that were sampled just 31
(0.61%) were inferred to be feral dogs (Table 3). Similarly, there
were only 27 animals identified as likely F1 or F2 dingo dog
hybrids. Contrary to widespread understanding, our results show
that feral dogs and feral dog dingo hybrids were very rare
across mainland Australia. This suggests that feral dogs have not
established a self-sustaining population in the wild and that inter-
breeding between dingoes and dogs may occur infrequently.
Our finding that feral dogs were rare and are unlikely to have
established a self-sustaining population on mainland Australia is
backed up by the rarity of true feral dog populations globally. In
Tasmania, there is a similar mix of European derived dog breeds
to mainland Australia and a similar environment to southeastern
Australia but there is little evidence that a feral dog population
has become established (DPIPWE 2013). The rarity of true feral
dogs both globally and in Australia suggests that domestic dogs
have not retained the ability to persist in the wild in the absence
of anthropogenic derived resources. Indeed, the Gala´ pagos
remains an isolated exemplar of domestic dogs establishing a
feral population (Barnett 1986;Reponen et al. 2014). However,
free breeding (‘village or camp dogs’) that rely on anthropogenic
food and water sources are of widespread occurrence in many
regions of the world (Gompper 2013;Home et al. 2018).
In Australia, free-breeding dogs are largely restricted to the
fringes of Indigenous communities (Collins and Mills 2013;
Newsome et al. 2013,2014;Hudson et al. 2018;Brookes et al.
2020;Ma et al. 2020).
Our collation of DNA ancestry testing results suggests that
most wild canids in Australia are pure dingoes (Fig. 1). Dog
introgression is uncommon in Western Australia, the Northern
Territory and South Australia, with more than of 90% of dingoes
tested in those states being pure dingoes. Of the dingoes which did
show domestic dog introgression, most carried more than 75%
dingo ancestry. No F1 or F2 dingo dog hybrids were observed in
the Northern Territory or South Australia. In Queensland, 68.5%
of wild canids were pure dingoes and a majority of the remaining
carried greater than 75% dingo ancestry. Only 7% of wild canids
in Queensland were less than 65% dingo, and 0.8% were feral
dogs or possible F1 or F2 dingo domestic dog hybrids.
As highlighted by Stephens et al. (2015), dog introgression is
most widespread in southeastern Australia (Fig. 2). Despite this,
the occurrence of possible F1 or F2 dingo dog hybrids was low
in southeastern Australia, making up less than 2% of the total
population inthese regions.It may be that dingo dog hybridisa-
tion events are rare in the wild or that the survival of wild canids
with less than 50% dingo ancestry is poor. The widespread
introgression of dog genes in southeastern Australia may reflect
backcrossing of F1 dingo dog hybrids, facilitating the spread of
dog genes into the wider dingo population over long periods of
time rather than a high occurrence of dingo dog hybridisation in
the wild. As emphasised by Stephens et al. (2015), dog introgres-
sion in dingoes may be more common in southeastern Australia
due to the earlier and more intensive European settlement,
resulting in increased contact between domestic dogs and dingoes
in these regions. Cairns et al. (2019) added that the widespread
occurrence of intensive lethal control, particularly aerial baiting,
may increase the likelihood of dingo dog hybridisation by
(a)
Pure dingo
Dingo (>75% ancestry)
Dingo (65–75% ancestry)
Dingo (50–65% ancestry)
No samples present
0 250 500 750 1000 km 0 250 500 750 1000 km
Feral dog (no dingo ancestry)
Feral dog (possible F1 dingo ×
dog hybrid)
Feral dog (no dingo ancestry)
Feral dog (possible F1 dingo ×
dog hybrid)
(b)
Fig. 2. Spatial patterns of dingo ancestry across Australia based on microsatellite DNA testing. (a) Median ancestry calculated for all samples within
each hex grid. As the results were consistent between mean and median ancestry, the map depicting mean ancestry is presented in Supplementary
Material S2. (b) Distribution of 31 feral domestic dogs and 27 feral dog hybrids (possible F1 or F2 dingo dog hybrids) across Australia. The positions
of the national dingo exclusion fence and the Western Australian dingo exclusion fence are depicted by solid black lines.
The wild dog myth Australian Mammalogy E

fracturing dingo social structures. Although this admixture from
dogs into the dingo population is a concern, it is important to note
that the dingo population still maintains a genetically and
morphologically dingo dominantidentity (Parr et al. 2016;Cairns
et al. 2019;Crowther et al. 2020).
There are several key knowledge gaps about the identity of
dingoes in Australia that bear consideration. First, we still lack
information about the genetic identity of dingoes across large
regions of Australia, particularly central and northern Australia
(Fig. 2). Morphological research about the phenotype of dingoes
with low levels of dog ancestry may assist on-ground manage-
ment and conservation efforts, particularly if distinguishing
features could be identified. Management plans should be
careful not to assume that a given population does or does not
carry domestic dog ancestry without the necessary genetic data.
Despite this, the broad pattern of dingo ancestry across Australia
suggests that in western, central and northern Australia, dog
introgression is likely to be limited and feral dogs extremely
rare. There is some concern that current microsatellite testing
methods may be biased by regional genetic variation within
dingoes (Cairns et al. 2017,2019). It is important to consider that
ancestry testing methods rely on the assumption that dingoes
form a single homogeneous population (Elledge et al. 2008;
Stephens et al. 2015;Cairns et al. 2019), an assumption we now
know to be false (Cairns and Wilton 2016;Cairns et al. 2018;
Koungoulos 2020). Robust dingo ancestry assessments require
broad sampling across Australia to capture regional genetic
variation (Cairns et al. 2019). Possibly some dingoes are mis-
classified as hybrids because of regional variation. As argued by
Cairns et al. (2019), the type and number of genetic markers
limits accuracy of genetic testing and estimates based on 23
microsatellites may not reflect genome-wide ancestry. Genome-
wide SNP genotyping may offer a cost-effective and high-
throughput alternative to address the limitations of microsatel-
lite genetic testing in the future. Thus, we caution managers and
researchers to evaluate the reliability of ancestry estimates and
urge end-users to explore technology improvements for ancestry
testing into the future.
There has been ongoing debate about the appropriate termi-
nology for wild canids in Australia, i.e. dingo or wild dog (Letnic
2012). Kreplins et al. (2019) found that the term wild dog was
more commonly used in studies funded by livestock industry
organisations, compared to conservation-based studies which
predominately used the term dingo. van Eeden et al. (2020)
studied public understanding of the terms dingo and wild dog.
They found that only 19.1% of respondents were aware that wild
dog control programs targeted dingoes and furthermore respon-
dents were generally not supportive of lethal dingo manage-
ment. At the 2019 Royal Zoological Society of NSW
symposium titled ‘Dingo Dilemma’ there was strong opposition
to the term wild dog, with many participants asserting that the
term wild dog disguises lethal management practices on dingoes
from the public and hinders debate about dingo management in
Australia (Dickman 2019). We add that the term wild dog does
not accurately represent the ancestry of wild canids in Australia,
particularly as the dominant genetic identity is dingo and feral
domestic dogs are virtually absent from the landscape (Fig. 2).
Although there are dingoes carrying domestic dog ancestry,
particularly in southeastern Australia, there are few F1 or F2
hybrids. The term hybrid generally refers to only F1 crosses, i.e.
the offspring of a dingo and domestic dog but F2 animals which
are the offspring of two F1 hybrids may also be referred to as
hybrids (Hansson et al. 2012). We suggest that dingoes that carry
domestic dog ancestry but are not F1/F2 hybrids should be
referred to as dingo backcrosses or admixed dingoes.
The finding that feral dogs have not established populations
has implications for the management of wild canids in Australia.
Dingoes and stray or roaming domestic dogs can cause serious
impacts for livestock graziers (Fleming et al. 2001). Management
of feral, stray or roaming domestic dogs should focus on
responsible pet ownership including spaying and neutering of
pet animals, keeping pet and working dogs under control and
confined during the night. As feral dogs do not represent a
significant portion of the wild canid population, it should be
clear in legislation and policy that lethal control programs are
targeting dingoes (including admixed dingoes) rather than feral
dogs. Although hybridisation is a concern in southeastern Aus-
tralia (Stephens et al. 2015;Cairns et al. 2019), responsible pet
ownership and continued exclusion of domestic dogs from
National Parks and conservation areas can reduce the occurrence
of future dingo dog hybridisation events. The low number of
F1 or F2 hybrids detected indicates that dingo dog hybridisa-
tion events are uncommon. Despite historical domestic dog
introgression, the dingo population maintains a dingo dominant
identity, even in southeastern Australia (Fig. 1). It is possible that
widespread lethal control programs have increased the likelihood
of dingo dog hybridisation events and facilitating the spread of
introgressed dog genes into the wider dingo population. Lethal
control has been identified as a factor increasing the likelihood of
interspecific hybridisation in other wild canids including coyotes
and red wolves by fracturing social structures and altering
demographic patterns (Bohling and Waits 2015). Management
programs that maintain stable dingo social structures present a
better balance to managing the risks to stock predation and dingo
conservation (Allen 2014,2015;Wallach et al. 2017). Addition-
ally, lethal control programs should not occur during the dingo
breeding season (winter) as this may facilitate dingo dog
hybridisation events, by fracturing pack structures and reducing
the availability of dingo mates. Baiting has also been linked to an
increase in the body-size of dingoes, possibly increasing their
impact on livestock (Letnic and Crowther 2020).
The lack of public engagement and debate on dingo conser-
vation on private and public lands in Australia has allowed
agricultural industry priorities to dominate government policy
and decision making on dingo management. Social science
studies show the general public are largely unawareof the current
threats wild dog control programs have on the remaining dingo
populations in Australia (van Eeden et al. 2020). The lack of
engagement by the public on dingo conservation can in part be
attributed to the renaming of the Australian dingo as a wild dog in
government literature and allowing the general misunderstanding
that all wild dogs are feral dogs to persist. We propose that a
terminology shift is required to reflect the identity of wild canids
in Australia: the term dingo needs to be reinstatedbecause genetic
testing demonstrates that a majority of animals are of high dingo
ancestry and feral dogs are virtually absent. The term wild dog
does not reflect the ancestry of wild canids in Australia and is
poorly understood by the public, it should be retired from use.
FAustralian Mammalogy K. M. Cairns et al.

Conflicts of interest
KMC is a scientific advisor to the Australia Dingo Foundation,
New Guinea Singing Dog Conservation Society and New
Guinea Highland Wild Dog Foundation.
Declaration of funding
KMC is supported by research funding from the Australian
Dingo Foundation.
Acknowledgements
The authors thank Barry Traill, Angus Emmott and David Pollock for the
impetus to write this paper following discussions at the RZS Dingo Dilemma
Symposium in 2019. We acknowledge the extensive work done by the late
Associate Professor Alan Wilton on this topic. The authors would also like to
thank two anonymous reviewers and the editor for their careful reading of
our manuscript and insightful comments that have improved this manuscript.
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The wild dog myth Australian Mammalogy I