Pelage variation in dingoes across southeastern Australia: implications for conservation and management

Abstract

How to manage hybridization and introgression in wild animals is controversial. Wildlife managers and researchers may often rely upon phenotypic variables such as coat colour to inform on ground management decisions. In Australia, dingoes are typically believed to be ginger in colour, and unusual coat colours such as brin-dle or sable are widely posited to be evidence of contemporary domestic dog hybridization. We carried out microsatellite-based genotyping on 1325 wild canids from southeastern Australia of known coat colour to estimate the extent of domestic dog introgression. A key aim of our study was to examine the relationship between coat colour and ancestry in wild dingoes. We observed that 27.4% of our samples were dingoes with no evidence of domestic dog ancestry whilst 72.6% were dingoes with some domestic dog ancestry. Our data confirm that feral dogs, domestic dogs with no dingo ancestry, are rare in the wild, representing less than 1.5% of the population. There was no coat colour that could distinguish dingoes with or without dog ancestry from each other. Contrary to popular belief, colours such as brindle and patchy were positively associated with dingoes with no dog ancestry and were less common in dingoes of mixed ancestry. A key finding of this work is that coat colour should not be used to assess ancestry in dingoes. Further research is needed to uncover the antiquity, origin and potential adaptive value of these genomic regions. It is possible that this is a similar example of adaptive introgression as has been observed in North American wolves with black coat colour. These data add perspective to global debates about how to manage and conserve enigmatic animal populations in the presence of modern or historical introgression.

Full text

Pelage variation in dingoes across southeastern Australia:
implications for conservation and management
K. M. Cairns
1,2
, K. D. Newman
3
, M. S. Crowther
4
& M. Letnic
1,2
1
Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW,
Australia
2
Evolution and Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney,
NSW, Australia
3
School of Biosciences, University of Melbourne, Parkville, VIC, Australia
4
School of Life and Environmental Sciences, University of Sydney, Sydney, NSW, Australia
Keywords
admixture; coat colour; dingo; dog; hybridization;
introgression; microsatellites; pelage.
Correspondence
Kylie M Cairns, Centre for Ecosystem Science,
School of Biological, Earth and Environmental
Sciences, University of New South Wales,
Sydney, NSW 2052, Australia.
Email: kylie@kyliecairns.com or
k.cairns@unsw.edu.au
Editor: Andrew Kitchener
Received 30 April 2020; revised 7 January 2021;
accepted 13 January 2021
doi:10.1111/jzo.12875
Abstract
How to manage hybridization and introgression in wild animals is controversial.
Wildlife managers and researchers may often rely upon phenotypic variables such
as coat colour to inform on ground management decisions. In Australia, dingoes
are typically believed to be ginger in colour, and unusual coat colours such as brin-
dle or sable are widely posited to be evidence of contemporary domestic dog
hybridization. We carried out microsatellite-based genotyping on 1325 wild canids
from southeastern Australia of known coat colour to estimate the extent of domes-
tic dog introgression. A key aim of our study was to examine the relationship
between coat colour and ancestry in wild dingoes. We observed that 27.4% of our
samples were dingoes with no evidence of domestic dog ancestry whilst 72.6%
were dingoes with some domestic dog ancestry. Our data confirm that feral dogs,
domestic dogs with no dingo ancestry, are rare in the wild, representing less than
1.5% of the population. There was no coat colour that could distinguish dingoes
with or without dog ancestry from each other. Contrary to popular belief, colours
such as brindle and patchy were positively associated with dingoes with no dog
ancestry and were less common in dingoes of mixed ancestry. A key finding of
this work is that coat colour should not be used to assess ancestry in dingoes. Fur-
ther research is needed to uncover the antiquity, origin and potential adaptive value
of these genomic regions. It is possible that this is a similar example of adaptive
introgression as has been observed in North American wolves with black coat col-
our. These data add perspective to global debates about how to manage and con-
serve enigmatic animal populations in the presence of modern or historical
introgression.
Introduction
Hybridization is a threat to many wildlife species and often
arises due to human modification of animal distributions allow-
ing previously allopatric species, sub-species or populations to
interbreed (Simberloff, 1996; Rhymer & Simberloff, 1996;
Allendorf et al., 2001). This has led to debate about the role
of adaptive introgression, blurring species boundaries and how
introgression should be managed for conservation (vonHoldt
et al., 2018; Supple & Shapiro, 2018; Muhlfeld et al., 2014;
Murphy et al., 2018; Mable, 2019; Macdonald et al. 2010).
Genetic introgression resulting from hybridization can be a dri-
ver of phenotypic variation (Anderson et al., 2009; Jones
et al., 2018; Schweizer et al., 2018; vonHoldt & Aardema,
2020). Consequently, phenotypic characteristics are often used
in management programmes to identify hybrids from non-hy-
brids (Kitchener et al., 2005; Daniels et al., 1998; Macdonald
et al. 2010; Elledge et al., 2006; Corbett, 2001a; Fleming
et al. 2001; Galov et al., 2015). However, discriminating
hybrid and non-hybrid forms on the basis of their phenotype
requires a sound understanding of the range of phenotypic
variability possible in both forms.
Dingo-dog hybridization is a controversial topic in Australia.
The domestication status of the dingoes’ancestor is unclear
and has sparked debate about the taxonomic status of dingoes.
Crowther et al. (2014) and Smith et al. (2019) suggest that
dingoes represent an early or pre-domestication dog lineage
that is distinct from modern domestic dogs, positing the
nomenclature Canis dingo. Jackson et al. (2017, 2019) argue
that dingoes are a feral domestic dog (Canis familiaris).
104 Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London
Journal of Zoology. Print ISSN 0952-8369

Regardless of taxonomy, dingoes are a wild living canine that
has been present in Australia for at least 3500–8000 years
according to molecular and archaeological evidence (Cairns &
Wilton, 2016; Fillios & Tac
ßon, 2016; Balme, O’Connor & Fal-
lon, 2018; Zhang et al., 2020). Since arriving on the continent,
dingoes have been subject to natural selection and display
many distinctive phenotypic traits (Crowther et al., 2014;
Smith et al., 2019; Zhang et al., 2020). However, dingoes can
hybridize with domestic dogs, and since British colonization of
Australia, there has been opportunity for dingoes to interbreed
with dogs (Newsome & Corbett, 1985; Stephens et al., 2015;
Cairns et al., 2019). This is particularly the case in eastern
Australia where opportunities for mating between dingoes and
dogs have been facilitated by higher densities of pet dogs and
active suppression of dingo populations (Stephens et al., 2015;
Cairns et al., 2019).
There has long been debate about the physical appearance
of dingoes and influence of hybridization on coat colour varia-
tion and skull morphology(Corbett, 2001b; Crowther et al.,
2014, Fleming et al. 2001; Elledge et al., 2006; Newsome &
Corbett, 1985; Jones, 1921; Macintosh, 1975; Barker & Macin-
tosh, 1979). Skull morphology was once widely used as a
method of discriminating between dingoes and hybrids (New-
some, Corbett & Carpenter, 1980; Corbett, 2001a; Elledge
et al., 2008). However, a recent study has shown that skull
morphology is highly conserved and may have limited utility
in discrimination of pure dingoes (Parr et al., 2016).
Historical reports from the 18th and 19th centuries com-
monly described observations of dingoes that were red, yellow,
black, white, black and white, tan or tawny (Collins, 1798;
Mitchell, 1839; Abbott, 2008). Whilst ginger (Fig. 1) is the
most common pelage colouration observed in genetically or
morphologically identified pure dingoes, other accepted colours
include black & tan and white (Elledge et al., 2006, Fleming
et al. 2001; Corbett, 2001a; Corbett, 2001b; Newsome & Cor-
bett, 1985; Crowther et al., 2014; Smith, 2015; Newsome
et al., 2013). Many dingoes, regardless of colour, have white
markings on their chest, feet, legs and tail tips, dark eyes and
undercoat that is white, cream or grey (Table 1). Sable (Fig. 1)
pelage has long been viewed as an indicator of dog gene intro-
gression by some authors (Fleming et al. 2001; Corbett,
2001b), but because it has been recorded in dingo pelts from
the early 1800s in remote regions it is likely to be another
‘wild-type’dingo colouration (Crowther et al., 2014). The
observation of pure sable dingoes by Tatler et al. (2021) in
remote South Australia provides further support that sable is
an ancestral colour variation. Other colour patterns (Fig. 1) that
have been attributed to be evidence of significant dog ancestry
include brindle, patchy (or parti colour), merle and brown
(Corbett, 2001b, Crowther et al., 2014, Elledge et al., 2006,
Fleming et al. 2001; Newsome & Corbett, 1985; Smith, 2015).
In the field, wild dog and dingo trappers often rely upon
coat colour to assess the likelihood of a dingo having domestic
dog ancestry (Elledge et al., 2008; Elledge et al., 2006; Flem-
ing et al. 2001; Fleming, Allen & Ballard, 2012). Some wild-
life managers base field assessments on the strict definition of
a dingo provided by Corbett (2001) which allows only animals
with ginger coat colour to be pure dingoes. Elledge et al.
(2008) found that visual diagnosis of wild dingoes by trappers
and wildlife managers based on their pelage characteristics was
not consistent with diagnoses derived from genetic methods.
Specifically, they found that their survey respondents only
defined dingoes with ginger or black & tan coat colour as pure
and that the ginger animals visually identified as pure were not
the same individuals identified by skull morphology or genetic
methods. This is concerning because it implies that visual
assessment of ancestry by trappers and wildlife managers may
not be accurate and that they may preferentially remove ‘non-
ginger’wild dingoes with unknown consequences for the
genetic integrity of populations. Indeed, there is widespread
appreciation that selective culling based on a specific pheno-
typic trait can influence the physical (and genetic) characteris-
tics of wild populations in unknown ways (Harris, Wall &
Allendorf 2002; Garel et al., 2007).
Another consideration is that some coat colour patterns may
have introgressed into the dingo population through hybridiza-
tion but have been maintained as they are neutral or confer
some selective advantage. In wolves (Canis lupus), the black
coat colouration introgressed from domestic dogs approxi-
mately 1500–7200 years BP (Anderson et al., 2009; Schweizer
et al., 2018). It is believed that once it entered the wolf gene
pool, the associated genetic regions conferred some advantage,
in the form of increased immunity, leading to its rapid spread
throughout the North America wolf population (Schweizer
et al., 2018). It is important to note here that black wolves are
still considered to be wolves not wolf-dog hybrids. This raises
the question of whether coat colour is an effective and useful
tool for field assessment of dingoes in Australia, particularly if
coat colours might reflect historical introgression.
Here we present coat colour and DNA purity assessment of
1325 wild canids from New South Wales and Victoria. We use
these data to examine the utility and accuracy of coat colour
in field-based ancestry assessments of dingoes, particularly in
southeastern Australia. These data also provide further knowl-
edge about the genetic and physical identity of dingoes in Aus-
tralia.
Materials and methods
Dingo samples
We obtained tissue samples from wild canids killed or trapped
through routine ‘wild dog’management activities in New
South Wales, The Australian Capital Territory and Victoria.
DNA of these individuals was sampled through either blood,
ear clips or buccal swabs. These samples were collected on
public and private lands between 1998 and 2014 and were
sent to UNSW (Alan N. Wilton and KMC) for a long-term
research programme on the ancestry of wild canids in Aus-
tralia. Upon arrival at UNSW, DNA was extracted using Qia-
gen DNeasy kits (Qiagen Sciences, Germantown, MD, USA)
and samples were genotyped using a widely used DNA test
for estimating admixture in dingoes (Wilton, Steward &
Zafiris, 1999; Wilton, 2001; Elledge et al., 2008; Stephens
et al., 2015; Cairns et al., 2019). The 23 microsatellites geno-
typed are distributed across the genome (Fig. 2). The data
Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London 105
K. M. Cairns et al. What colour can a dingo be?

used in this study come from a large database of ancestry esti-
mates and biological data from over 4000 wild and captive
dingoes across Australia.
For the purpose of this study, we identified wild samples
which had coat colour information recorded, this resulted in
1325 samples with raw genotype data from more than 14
microsatellites (out of the total 23) and coat colour information
(Table S1). We restricted the samples to southeastern Australia
as this is where a majority of the dingo samples came from
and we did not wish to introduce geographic bias which may
be present between southeastern and northwestern dingo popu-
lations (Cairns & Wilton, 2016; Cairns et al., 2017; Cairns
et al., 2018).
Ancestry analysis
We carried out ancestry estimation using the STRUCTURE
method as per Cairns et al. (2019). Briefly, simulations were
run in STRUCTURE v2.3.4 (Pritchard, Stephens & Donnelly,
2000; Falush, Stephens & Pritchard, 2003) using the 1325 wild
canids and a set of pre-defined reference populations. We used
the extended reference population with 50 dingoes and 66 dogs
to allow for geographic variation in ancestry estimates from
Cairns et al. (2019). The dog reference population contains
mixed breed dogs to provide a genetically diverse domestic
dog population compared to dingoes, with the knowledge that
dingoes form a distinct evolutionary lineage compared to
domestic dogs (Cairns et al., 2018).
Analyses were run in STRUCTURE (Pritchard, Stephens &
Donnelly, 2000; Falush, Stephens & Pritchard, 2003) with the
admixture and correlated allele frequency models, default set-
tings for alpha were used. As K=2 was identified as the most
appropriate Kfor these data (Stephens et al., 2015; Cairns
et al., 2019), we ran simulations for only K=2 for 200 000
iterations with a 20 000 iteration burn-in period, and 10 repli-
cates were performed. All simulations were run with the USE-
POPFLAG on. CLUMPAK was used to average individual q
values across the 10 replicates (Kopelman et al., 2015). We
FIGURE 1 A selection of photographs of wild dingoes from Kosciuszko National Park in southeastern Australia. Examples of sable (a), ginger (b),
black & tan (c), brindle (d) and patchy (e) coat colour patterns are evident. See Table 1 for a detailed description of coat colour patterns.
Photographs courtesy of Michele J Photography, Cooma NSW. [Colour figure can be viewed at zslpublications.onlinelibrary.wiley.com.]
106 Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London
What colour can a dingo be? K. M. Cairns et al.

then assigned individuals to categories based on their average
q value representing dingo ancestry (Cairns et al., 2019).
Pelage variation in wild canids
Coat colour descriptions provided by sample collectors on sub-
mission were categorized into the following fields: ginger,
white, sable, black & tan (or white) points, brindle, patchy and
black. Analysis was carried out in SPSS v26 (IBM, Armonk,
NY) and R v3.6.2 (R Development Core Team, 2010) with
packages ‘graphics’and ‘corrplot’using the dataset of 1325
animals with records for coat colour and genetic ancestry esti-
mates. For statistical analyses, we categorized the animals as
follows: dingoes with only dingo ancestry, that is pure dingoes
(q value >0.8); dingoes with greater than 75% ancestry (q
value between 0.7 and 0.79) and dingoes with 75–50% ances-
try (q value between 0.5–0.69) based on Cairns et al. (2019)
and Stephens et al. (2015). First, we summarized observed
coat colour between the three categories of dingoes in SPSS
v26. Then, in R v3.6.2, we performed Pearson’s chi-squared
tests of independence to compare coat colours between dingoes
and dingoes with varying degrees of dog ancestry. We calcu-
lated adjusted standardized residuals (ASR’s) from the contin-
gency table to investigate where differences in coat colour
were observed. ASR’s of greater than 2 or less than 2 indi-
cate departure from the null hypothesis.
Results
Ancestry analysis
According to the STRUCTURE simulations using extended
Wilton dingo and dog reference populations, 27.4 % of the
1325 samples were classified as having only dingo ancestry or
were likely to only have dingo ancestry (q value >0.8). 40.3
% of samples were admixed dingoes with greater than 75%
dingo ancestry (q value 0.7–0.79) and 32.3 % of samples were
dingoes with between 50 and 75% dingo ancestry (Fig. 3, Fig-
ure S1, Table S1). There were only 5 feral domestic dogs
(0.4%) and 15 feral domestic dog hybrids with less than 50%
dingo ancestry (1.1%) out of 1325 wild dingo samples; they
were removed from subsequent analyses due to the small sam-
ple size (Fig. 3, Figure S1, Table S1).
Pelage variation in wild canids
Dingoes in our sample set displayed a variety of coat colours
(Table 2, Table S1). Across our 1305 dingo samples, we
Table 1 Description of commonly observed dingo (Canis dingo) coat colour patterns
Coat Colour Pattern Description
Ginger A solid red coat varying in shade from deep red to sandy yellow to warm cream. Occasional interspersed
black hairs present on muzzle which may be dark in juvenile and pale out to grey by puberty; and
sometimes appear on sides of neck and shoulder region or top of tail. Often present are white markings
particularly on the chest, feet, legs and tail tip. Often, the colouration is lighter on the underbelly.
Undercoat where present varies from white to grey. The tail may exhibit cream and/ or agouti
undercoat. Eyes dark.
White A solid whitish or pale cream coat with no black pigment throughout. Can also exhibit white spotting on
chest, neck, digits and extremities Eye rims remain pigmented. Nose can pale to liver colour in winter.
Undercoat when present is white. Iris pigment remains dark.
Black & tan A solid black coat with tan points, brow pips, muzzle, cheek spots, chest, belly, feet and legs. Lighter
underbelly. The markings may vary from deep tan to cream, also with white spotting at extremities. The
undercoat, if present, is pale cream or grey. The tail may show both cream and grey undercoat. Eyes
dark brown.
Black A solid black coat. May be white markings particularly on the chest, feet, legs and tail tip. The undercoat
when present is dark grey. Eyes dark.
Sable A ginger to cream base coat with dark juvenile muzzle, greying to adult form. Black tipped hairs spreading
from the midline and interspersing down shoulder area and along upper line of tail. White extremities,
tail tip and chest markings. feet, etc. In some cases, the coat colour may appear almost dark grey or
there may be a distinct saddle pattern along the back. Underbelly lighter in colour. Undercoat may be
white, cream or greyish. Eyes dark brown.
Brindle A coat pattern with ginger background and irregular dark banding across midline spreading down sides of
body and legs. May also exhibit white extremities and chest markings. Undercoat where present may
be white or cream. Eyes dark.
Patchy (parti colour) A basic ginger or sandy or black and tan base coat pattern broken with extensive white markings, forming
patches. The white markings may run together and be so extensive as to leave only small areas of
ginger or black ie on the body or head, or form a white-collar pattern. White extremities and tail tip.
Undercoat where present is white. Eyes dark brown.
Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London 107
K. M. Cairns et al. What colour can a dingo be?

observed the following coat colours: 53.5% ginger, 10.7%
black & tan, 5.5% black, 1.4% white, 13.9% brindle, 8.9%
sable and 6.2% patchy (Table 2). All of the coat colours were
observed in dingoes and dingoes with varying degrees of dog
ancestry (Fig. 4). Comparing the proportion of colours
observed in dingoes (q value >0.8), dingoes with greater than
75% dingo ancestry (q value 0.7–0.79) and dingoes with
between 50 and 75% dingo ancestry (q value 0.5–0.69), we
observe that ginger is the most common colour of all dingoes
(Fig. 4). We observed feral dogs with ginger, black & tan and
sable coat colours (Table 3).
There are significant associations between coat colours and
their genetic ancestry (v
2
=58.052, d.f. =12, P-
value <0.001). We used ASR’s to identify where the differ-
ences in coat colour between dingo categories occurred
(Table 4, Fig. 5). There was a strong positive association
between pure dingoes and the colour brindle (ASR =3.31)
and a strong negative association between pure dingoes and
the colour black (ASR =2.42). There was a weakly negative
association between pure dingoes and the colour black & tan
(ASR =1.96). There was a strong positive association
between 50 and 75% dingoes and the coat colours black
(ASR =2.24) or black & tan (ASR =2.56), but a very strong
negative association with the coat colours brindle
(ASR =3.19; Fig. 5). There was a weak negative association
between 50–75% dingoes and the colours patchy
(ASR =1.98) and White (ASR =1.99). The strongest dif-
ferences between dingo categories were based on the colours
brindle, black and black & tan (Fig. 5). Dingoes with >75%
ancestry had no positive or negative associations with any of
the pelage colours, indicating that they were not more or less
likely to carry any of the coat colourations.
Discussion
Coat colour is highly variable in wild dingoes across southeast-
ern Australia (Newsome & Corbett, 1985; Corbett, 2001b) and
our data show it is not a reliable measure of domestic dog
introgression. We observed pure dingoes with coat colour pat-
terns such as brindle and patchy (Fig. 4), colours often
asserted as being observed only in dingo-dog hybrids or dogs
(Corbett, 2001b; Elledge et al., 2006; Fleming et al. 2001;
Smith, 2015). There were differences in the proportion of coat
colour patterns between pure dingoes, dingoes with low levels
of dog ancestry and dingoes with moderate dog ancestry
(Fig. 4). However, there was no coat colour that could be used
to distinguish between pure dingoes and those with dog intro-
gression (Fig. 4). Therefore, we caution wildlife managers and
conservation organizations against using coat colour to assess
dog introgression in dingo populations.
Our findings indicate there are some differences in the fre-
quency of coat colours between pure dingoes and those carry-
ing dog ancestry (Figs. 4 and 5, Table 4). For example, whilst
white coat colour was observed in pure dingoes and those with
>75% dingo ancestry, it was very rare in dingoes with 50-75%
dingo ancestry. Surprisingly, patchy coat colour was most
Figure 2 Chromosome and relative position of the 23 microsatellite markers used in dingo DNA ancestry testing and the coat colour genes
ASIP,RALY,MITF,MC1R and CBD103 which are responsible for coat colour variation in canines [Colour figure can be viewed at zslpublications.
onlinelibrary.wiley.com.]
108 Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London
What colour can a dingo be? K. M. Cairns et al.

commonly observed in pure or high ancestry dingoes (Fig. 4),
contradicting the widespread belief that animals with patchy
coats are likely to be hybrids (Corbett, 2001b; Elledge et al.,
2006, Fleming et al. 2001; Newsome & Corbett, 1985). Brin-
dle, a coat colour, commonly cited to be a result of dog
hybridization (Corbett, 2001b; Elledge et al., 2006; Fleming
et al. 2001), was strongly associated with pure dingoes and
negatively associated with dingoes carrying 25–50% dog
ancestry (Fig. 5). This suggests that brindle colouration is not
informative for distinguishing dingoes with or without dog
introgression. Whilst black or black & tan colours were posi-
tively associated with lower ancestry dingoes (20–25% dog
ancestry, Fig. 5), the black & tan colour pattern is well estab-
lished as being an ancestral dingo colour, limiting the utility of
this result for wildlife managers. Sable was observed relatively
evenly between pure dingoes and those carrying dog ancestry,
suggesting limited association with hybridization (Fig. 4).
These data are consistent with observation of sable colouration
in contemporary pure dingo populations (Tatler et al., 2021) as
well as historical pelt collections (Crowther et al., 2014), sug-
gesting that sable colouration be considered an ancestral colour
variation and challenging its use as an indicator of domestic
dog introgression. The findings of this study challenge the
widespread understanding that coat colour can be used to iden-
tify hybrids (Corbett, 2001b; Elledge et al., 2006; Fleming
32.3%
Dingo (50-75%) 40.3%
Dingo (>75%)
27.4%
Pure Dingo
1.1%
Feral dog (<50%)
0.4%
Feral dog
}
} Feral dogs (Dogs + <50% dingo)
Dingoes (Pure dingoes + >50%)
Figure 3 Pie chart depicting proportion of pure dingoes, dingoes with
>75% ancestry, dingoes with 50-75% ancestry and feral dogs based
on STRUCTURE analysis of 23 microsatellites. [Colour figure can be
viewed at zslpublications.onlinelibrary.wiley.com.]
Table 2 Ancestry estimates of 1325 wild canids (Canis dingo or Canis familiaris) in southeastern Australia from STRUCTURE modelling of 23
microsatellites
Ancestry Structure q value range Number of individuals (n) Percentage
Dingo 0.8–1.0 358 27.4
Dingo with >75% dingo ancestry 0.7–0.79 526 40.3
Dingo with 50-75% dingo ancestry 0.5–0.69 421 32.3
Feral dog with <50% dingo ancestry 0.25–0.49 15 1.1
Feral dog with no dingo ancestry 0.24 5 0.4
Dingo with dog
ancestry
(50-75% dingo)
Dingo with dog
ancestry
(>75% dingo)
Dingo
Percent
100
80
60
40
20
0
White
Sable
Patchy
Ginger
Brindle
Black & Tan
Black
Coat Colours
Figure 4 Observed coat colour patterns in 1305 dingoes, dingoes with >75% ancestry and dingoes with 50–75% ancestry. [Colour figure can be
viewed at zslpublications.onlinelibrary.wiley.com.]
Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London 109
K. M. Cairns et al. What colour can a dingo be?

et al. 2001, Newsome & Corbett, 1985). Indeed, some coat
colours commonly believed to be ‘hybrid’colours were more
commonly observed in pure or high ancestry dingoes.
The most common colour for both dingoes and dingo
hybrids in our sample was ginger (ranging from cream to
sandy and red), consistent with observations of dingoes across
Australia (Newsome et al., 2013; Tatler et al.,2021; Newsome
& Corbett, 1985; Fleming et al. 2001; Corbett, 2001b).
Approximately 53% of the population exhibited this coloura-
tion, irrespective of ancestry (Table 2, Figure 4). This finding
is consistent with observational studies of the frequency of
pelage colouration in southeastern Australia. However, in
northern, western and central Australia 70-90% of the popula-
tion have ginger colouration (Newsome & Corbett, 1985; Cor-
bett, 2001b). Previous authors have commented that the greater
range of phenotypic variation in dingoes from southeastern
Australia was a result of higher levels of domestic dog
hybridization (Corbett, 2001b; Fleming et al. 2001; Newsome
& Corbett, 1985). However, recent genomic studies have
demonstrated that there are multiple evolutionary lineages of
dingo which are geographically subdivided (Cairns & Wilton,
2016; Cairns et al., 2017). Thus, it is possible that the south-
eastern and northwesterm lineages have different phenotypic
appearances unrelated to hybridization.
It is plausible that some coat colour patterns such as brindle
and patchy have originated from dingo-dog hybridization, and
that subsequent backcrossing has spread these colours into the
dingo population with little remnant dog ancestry remaining. It
Table 3 Observed coat colours of feral dogs (Canis familiaris)in
southeastern Australia across different categories of ancestry and
total observed proportion of coat colours across dataset
Feral dog
hybrid (n)
Feral
dog (n)
All feral dog
categories (%)
Black 0 1 5.3
Black & tan 3 0 15.8
Brindle 0 0 0.0
Ginger 10 2 63.2
Patchy 0 0 0.0
Sable 1 2 15.8
White 0 0 0.0
Table 4 Observed coat colours of dingoes (Canis dingo)in
southeastern Australia across different categories of ancestry and
total observed proportion of coat colours across dataset
Dingo (n)
Dingo >75%
(n)
Dingo
50–75% (n)
All dingo
categories (%)
Black 9 29 34 5.5
Black & tan 26 51 62 10.7
Brindle 73 74 34 13.9
Ginger 186 280 232 53.5
Patchy 30 35 16 6.2
Sable 26 48 42 8.9
White 8 9 1 1.4
Black
Black and Tan
Brindle
Ginger
Patchy
Sable
White
Dingo
Dingo with dog ancestry
(>75% dingo)
Dingo with dog ancestry
(50-75% dingo)
Standardized
Residuals:
<-4 -4:-2 -2:0 0:2 2:4 >4
-2.42
-1.85
-1.27
-0.7
-0.13
0.45
1.02
1.59
2.17
2.74
3.31
Black
Black and Tan
Brindle
Ginger
Patchy
Sable
White
Dingo
Dingo with
dog ancestry
(>75% dingo)
Dingo with
dog ancestry
(50-75% dingo)
0
1.89
3.78
5.67
7.56
9.45
11.35
13.24
15.13
17.02
18.91
Dingo
Dingo with
dog ancestry
(>75% dingo)
Dingo with
dog ancestry
(50-75% dingo)
Black
Black and Tan
Brindle
Ginger
Patchy
Sable
White
(a) (b) (c)
Figure 5 Adjusted standardized residual (ASR’s) analyses of coat colour variation between dingoes, dingoes with >75% ancestry and dingoes
with 50–75% ancestry. (a) Mosaic plot of ASR results, values of greater than 2 or less than 2 indicate departure from the null hypothesis. The
size of tiles reflects the sample size (n) within each category. (b) ASR correlation plot to investigate positive and negative associations between
rows and columns in the contingency analysis with positive values as blue circles and negative values as red circles. (c) ASR contribution plot
indicating the relative contribution of each cell to the contingency table analysis. [Colour figure can be viewed at zslpublications.onlinelibrary.wile
y.com.]
110 Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London
What colour can a dingo be? K. M. Cairns et al.

is also possible that coat colour variation in dingoes has some
adaptive potential. For example, introgression from Black-tailed
jackrabbits (Lepus californicus) into Snowshoe hares (Lepus
americanus) has led to adaptive variation in winter-moult col-
ours in areas with low snow cover (Jones et al., 2018). Black
coat colour in North American wolves is the result of intro-
gression from early American dogs approximately 1500–
7200 years ago (Anderson et al., 2009; Schweizer et al.,
2018). Schweizer et al. (2018) found that the melanistic coat
colour spread widely through the wolf population in a selective
sweep, because surrounding genomic regions conferred
enhanced immunity. Strikingly, the same gene (CBD103)is
responsible for solid black and brindle colouration (Kerns
et al., 2007; Dreger et al., 2019), plausibly the appearance of
brindle coat colour in dingoes may have some adaptive benefit
which warrants further investigation with genome-wide data.
Whilst it is possible that ‘non-typical’colours such as brin-
dle, patchy and sable are the result of introgression from dogs
into the dingo population, it is also possible that these colours
are ancestral to dingoes but were poorly described by early
European explorers. There are few historical records describing
dingo colours and those that exist, use only basic descriptors
such as ‘yellow-dog’(Abbott, 2008), ‘red with some white
about it’(Collins, 1798) or ‘black native dog’(Mitchell,
1839). A synthesis of museum records and historical accounts
of coat colour in dingoes suggest that at the time of European
arrival, dingoes were predominately ginger, sable, black and
tan, white or black (Jones, 1921; Macintosh, 1956; Macintosh,
1975; Elledge et al., 2006; Abbott, 2008; Crowther et al.,
2014); however, this cannot exclude the possibility that other
colours were present.
The antiquity of other pelage colourations such as patchy
and brindle is unclear. Macintosh recorded coat colour varia-
tion across much of Australia and observed that it was highly
variable with some regional differences (Macintosh, 1956;
Barker & Macintosh, 1979). There have been no verified
reports of brindle or patchy coat colour dingoes by early
explorers, implying these may not be ancestral (Troughton,
1958; Barker & Macintosh, 1979; Newsome & Corbett, 1985;
Crowther et al., 2014). More detailed genetic data such as
whole genome sequencing or interrogation of archaeological
remains will be needed to identify the antiquity these pelage
patterns in dingoes. Nevertheless, Newsome & Corbett (1985)
reported that 9% of their morphologically identified pure
dingo samples from southeastern Australia were brindle, sug-
gesting that its occurrence in wild dingoes has been relatively
stable over the last 60 years and its prevalence is unlikely to
be the result of contemporary dingo-dog introgression. The
origin of patchy coat colour may be more complicated. Din-
goes are often characterized as having white points such as
white feet, socks, toes and/or tail tips (Corbett, 2001b; Elledge
et al., 2006; Fleming et al. 2001) and genomically these
markings are controlled by the MITF gene (Schmutz, Berryere
& Dreger, 2009). According to Chew et al. (2019), some din-
goes may carry a MITF genotype predictive of extreme white
or piebald markings. Intriguingly, Macintosh (1956) observed
that patchy coat colour (ginger with white patches and/or col-
lars; Fig. 6) arose after multiple generations of inbreeding in a
captive bred colony. It is possible that the extent of white
markings in dingoes is controlled by MITF, a co-dominant
inheritance pattern and an unknown modifier gene (Karlsson
et al., 2007; Baranowska K€
orberg et al., 2014; Chew et al.,
2019), manifesting in dingoes carrying a range of phenotypes
between solid ginger with white points and patchy (Figs. 1
and 6). This might explain the appearance of excessive white
markings in some dingoes without the presence of dog intro-
gression. Future research should aim to use whole genome or
genome-wide SNP data to investigate the timing, genomic
mechanisms and possible origin of patchy and brindle coat
colours in dingoes.
There has been ongoing debate about the genetic identity of
dingoes and management of hybridization. In the wild, DNA
testing demonstrates that most dingoes are pure or backcrosses
with high levels of dingo ancestry and less than 1.1% of the
population were estimated to be less than 50% dingo ancestry
(Stephens et al., 2015; Cairns et al., 2019). Our data here
depict a population where 27.4% are pure dingoes, 40.3% were
>75% dingo and 32.3% were 50–75% dingo (Fig. 3,
Table S1). We observed 15 feral dog hybrids with less than
50% dingo ancestry and only 5 feral domestic dogs within our
dataset. Allen et al. (2017) suggest that a dingo should be con-
sidered pure if it is assessed as more than 93% dingo ancestry.
Others suggest that strict genetic thresholds are not useful, par-
ticularly if mixed ancestry dingoes are morphologically and
ecologically indistinguishable from pure dingoes (van Eeden
et al., 2018; Cairns et al., 2019). Indeed, tolerance of dingoes
with a small portion of dog genes (ie <25% dog ancestry) and
maintenance of stable dingo packs may limit future dingo-dog
hybridization, but this would require cessation of landscape
level aerial and ground baiting programmes (Miller, Adams &
Waits, 2003; Elledge et al., 2008; Cairns et al., 2019).
Genetic monitoring is useful for wildlife managers to
observe patterns of introgression across the landscape and time,
but as genetic estimates are not real-time DNA testing cur-
rently has limited utility to individual culling decisions. It is
also important for end-users of DNA testing to consider the
accuracy and reliability of ancestry estimates, for example
microsatellite testing may only be able to identify dog
hybridization within 4 generations (Cairns, Wilton & Ballard
2011). Data presented here indicate that coat colour also has
limited utility in field assessment of wild dingoes and may
lead wildlife managers to cull high ancestry dingoes. Indeed,
wildlife managers may be more likely to cull brindle dingoes
rather than ginger dingoes (Fleming et al. 2001, Fleming,
Allen & Ballard, 2012), but our data suggest that brindle
colouration is more actually common in pure dingoes than
hybrids (Figs. 4 and 5). Coat colour may appear to be an
innocuous trait, but it could have adaptive potential in dingoes,
as has been observed in wolves and other species (Hamilton &
Miller, 2016; Schweizer et al., 2018; Jones et al., 2018).
As our knowledge about the occurrence of interspecific intro-
gression and the adaptive potential of introgression increases, so
too does debate about how introgression is managed in wild pop-
ulations (Hamilton & Miller, 2016; Supple & Shapiro, 2018; von-
Holdt et al., 2018; Mable, 2019; vonHoldt & Aardema, 2020). In
North America and Europe, there is ongoing discussion of how to
Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London 111
K. M. Cairns et al. What colour can a dingo be?

manage and conserve canids despite the occurrence of admix-
ture between wolves, red wolves (Canis rufus), coyotes (Canis
latrans) and dogs (vonHoldt & Aardema, 2020, vonHoldt
et al., 2018, vonHoldt et al., 2016, Donfrancesco et al. 2019,
Hendricks et al., 2019). There is an important balance in pro-
moting the preservation of species genomic integrity, by limit-
ing future introgression but this may not include ‘reversing’
introgression or removing introgressed individuals from a pop-
ulation (Toro, Villanueva & Fern
andez, 2014; vonHoldt &
Aardema, 2020). In canids, ongoing research into recovery
programmes has identified that introgression is facilitated by
the disruption of social structures and low availability of con-
specific mates (Bohling & Waits, 2015; Bohling et al., 2016).
This is why many scientists are now arguing against intensive
suppression of dingo populations (Wallach et al., 2009; van
Eeden et al., 2018; Cairns et al., 2019), which likely increases
the risk of dingo-dog matings. We now add that wildlife man-
agers, conservation organizations and the public should not
rely upon coat colour to assess the ancestry of dingoes or
other wild canids.
Acknowledgments
The authors acknowledge the contributions of the late A/Prof
Alan Wilton (UNSW) who carried out much of the microsatel-
lite genotyping and pioneered genetic research on dingoes until
2011. Thank you to Ms Lyn Watson from the Australian
Dingo Foundation for discussions about coat colour in dingoes,
assistance with descriptive definitions of coat colours and
locating a photograph of NWG Macintosh with one of his pat-
chy dingoes.
Conflict of interest
KMC is a scientific advisor to the Australian Dingo Founda-
tion, New Guinea Highland Wild Dog Foundation and New
Guinea Singing Dog Conservation Society. KDN is a volunteer
at the Australian Dingo Foundation. No other interests
declared.
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Supporting Information
Additional Supporting Information may be found in the online
version of this article:
Fig S1. Ancestry coefficient estimates of 1325 wild dingoes
collected across southeastern Australia according to STRUC-
TURE analysis of 23 microsatellites. The dingo reference pop-
ulation (Dingo ref) contains 50 animals and the dog reference
population (Dog ref) includes 66 animals.
Table S1. Table of 1325 wild canid samples with coat colour
and STRUCTURE modelling ancestry estimate.
K. M. Cairns et al. What colour can a dingo be?
Journal of Zoology 314 (2021) 104–115 ª2021 The Zoological Society of London 115