An updated description of the Australian dingo (Canis dingo Meyer, 1793)

Abstract

A sound understanding of the taxonomy of threatened species is essential for setting conservation priorities and the development of management strategies. Hybridization is a threat to species conservation because it compromises the integrity of unique evolutionary lineages and can impair the ability of conservation managers to identify threatened taxa and achieve conservation targets. Australia's largest land predator, the dingo Canis dingo, is a controversial taxon that is threatened by hybridization. Since their arrival <5000 yBP (years Before Present) dingoes have been subject to isolation, leading to them becoming a unique canid. However, the dingo's taxonomic status is clouded by hybridization with modern domesticated dogs and confusion about how to distinguish ‘pure’ dingoes from dingo-dog hybrids. Confusion exists because there is no description or series of original specimens against which the identities of putative hybrid and ‘pure’ dingoes can be assessed. Current methods to classify dingoes have poor discriminatory abilities because natural variation within dingoes is poorly understood, and it is unknown if hybridization may have altered the genome of post-19th century reference specimens. Here we provide a description of the dingo based on pre-20th century specimens that are unlikely to have been influenced by hybridization. The dingo differs from the domestic dog by relatively larger palatal width, relatively longer rostrum, relatively shorter skull height and relatively wider top ridge of skull. A sample of 19th century dingo skins we examined suggests that there was considerable variability in the colour of dingoes and included various combinations of yellow, white, ginger and darker variations from tan to black. Although it remains difficult to provide consistent and clear diagnostic features, our study places morphological limits on what can be considered a dingo.

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An updated description of the Australian dingo
(Canis dingo Meyer, 1793)
M. S. Crowther1, M. Fillios2, N. Colman3& M. Letnic4
1 School of Biological Sciences, University of Sydney, Sydney, NSW, Australia
2 Department of Archaeology, University of Sydney, Sydney, NSW, Australia
3 Hawkesbury Institute for the Environment, University of Western Sydney, Penrith, NSW, Australia
4 Centre for Ecosystem Science, and School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW,
Australia
Keywords
apex predators; conservation; dogs;
hybridization; size; taxonomy.
Correspondence
Mathew Samuel Crowther, School of
Biological Sciences, University of Sydney,
Sydney, NSW 2006, Australia. Tel: +61 2
9351 7661; Fax: +61 2 9351 4119
Email: mathew.crowther@sydney.edu.au
Received 26 June 2013; revised 20
February 2014; accepted 25 February 2014
doi:10.1111/jzo.12134
Abstract
A sound understanding of the taxonomy of threatened species is essential for
setting conservation priorities and the development of management strategies.
Hybridization is a threat to species conservation because it compromises the
integrity of unique evolutionary lineages and can impair the ability of conserva-
tion managers to identify threatened taxa and achieve conservation targets.
Australia’s largest land predator, the dingo Canis dingo, is a controversial taxon
that is threatened by hybridization. Since their arrival <5000 yBP (years Before
Present) dingoes have been subject to isolation, leading to them becoming a
unique canid. However, the dingo’s taxonomic status is clouded by hybridization
with modern domesticated dogs and confusion about how to distinguish ‘pure’
dingoes from dingo-dog hybrids. Confusion exists because there is no description
or series of original specimens against which the identities of putative hybrid and
‘pure’ dingoes can be assessed. Current methods to classify dingoes have poor
discriminatory abilities because natural variation within dingoes is poorly under-
stood, and it is unknown if hybridization may have altered the genome of post-
19th century reference specimens. Here we provide a description of the dingo
based on pre-20th century specimens that are unlikely to have been influenced by
hybridization. The dingo differs from the domestic dog by relatively larger palatal
width, relatively longer rostrum, relatively shorter skull height and relatively wider
top ridge of skull. A sample of 19th century dingo skins we examined suggests that
there was considerable variability in the colour of dingoes and included various
combinations of yellow, white, ginger and darker variations from tan to black.
Although it remains difficult to provide consistent and clear diagnostic features,
our study places morphological limits on what can be considered a dingo.
Introduction
A sound understanding of the taxonomy of threatened taxa is
essential for setting conservation priorities and the develop-
ment of species management strategies (Mace, 2004). A poor
understanding of species taxonomy can hamper biodiversity
conservation efforts by preventing the identification of unique
evolutionary units, particularly if the species of potential con-
servation concern possesses morphological traits that are
similar to those of closely related species (Daugherty et al.,
1990). This is particularly true in canids where separate lin-
eages easily hybridize and produce fertile offspring (Roy et al.,
1994). Without the taxonomic tools to identify unique evolu-
tionary lineages, it may not be possible to make accurate
population estimates of species, identify threatened taxa or
develop management strategies to enhance the conservation
status of threatened taxa (Bacon & Bailey, 2006).
Australia’s largest land predator, the dingo (also known in
Australia as wild dog), is an example of a controversial taxon
that is threatened by hybridization with domestic dogs. Based
on molecular (Savolainen et al., 2004) and archaeological evi-
dence (Gollan, 1984), dingoes have been present on the Aus-
tralian continent for at least 3000–5000 years. Genetic
evidence suggests that dingoes originated from domestic dogs
from East Asia (Oskarsson et al., 2011). Since its arrival in
Australia and prior to the arrival of European colonists, the
dingo had been subject to at least 3000 years of isolation from
other canids, and presumably had been subject to genetic drift,
and natural selection, leading to it become a unique canid
(Corbett, 1995).
Recent research has documented the positive role that
dingoes have on biodiversity conservation through their regu-
lation of trophic cascades (Letnic, Ritchie & Dickman, 2012).
In particular, dingoes appear to benefit species threatened by
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invasive red foxes, owing to their suppressive effects on fox
abundance. However, efforts to harness the ecological inter-
actions of dingoes are hampered by the uncertain taxonomy of
the dingo (Letnic et al., 2012). In particular, the dingo’s taxo-
nomic status is clouded by hybridization with feral dogs and
confusion about how to distinguish ‘pure’ dingoes from
dingo-dog hybrids (Radford et al., 2012).
The poor taxonomic discrimination of dingoes from their
hybrids with feral dogs is of particular concern as dingoes and
dingo/dog hybrids are considered major pests to agriculture
because they kill livestock, and current policies in some juris-
dictions of Australia aim to exterminate dingo-dog hybrids,
but conserve dingoes (Letnic et al., 2012). Confusion exists, in
part, because the scientific description of Canis dingo (Kerr,
1792; Meyer, 1793) is based on a rudimentary picture (Fig. 1)
and brief description included in the journal of Australia’s first
colonial governor, Arthur Phillip (Mazell & Phillip, 1789),
and there is no surviving original specimen against which the
identities of putative hybrid and ‘pure’ dingoes can be
assessed. The dingo was first named as Canis antarticus (Kerr,
1792) based on the picture and description given by Arthur
Phillip (Mazell & Phillip, 1789). However, a subsequent
description of C. dingo based on the same material was given
by Meyer (1793). The name C. antarticus was suppressed in
favour of C. dingo because the latter name was in common
usage [International Commission of Zoological Nomencla-
ture (ICZN) 1957]. Since its initial description, other names
have been proposed for the dingo such as C. familiaris
australasiae (Desmarest, 1820), C. australiae (Gray, 1826),
C. dingoides (Matschie, 1915) and C. macdonnellensis
(Matschie, 1915).
Although the dingo has been subject to various reclassifi-
cations and changes in nomenclature, debate remains over
what morphological characters can be used to distinguish
dingoes, feral dogs and their hybrids (Jones, 2009; Radford
et al., 2012). Visual assessment of external characters is the
most common technique for classifying dingoes, feral dogs
and their hybrids. This approach relies upon the use of expert
knowledge to identify traits that distinguish dingoes from
hybrids. However, Elledge et al. (2008) found some corrobo-
ration between classifications made through genetic and skull
analysis methods, but none between either analytical method
and visual assessment. Similarly, Newsome & Corbett (1985)
could not distinguish between individuals classified using skull
measurements as dingoes or dingo-dog hybrids on the basis of
their coat coloration. Newsome, Corbett & Carpenter (1980)
and Newsome & Corbett (1982) used measurements of skull
morphology to discriminate dingo, dog and hybrid skulls, but
did not know the level of hybridization within the dingo
samples. Molecular studies that have attempted to discrimi-
nate between the genotypes of dingoes and their hybrids have
used captive animals held by breeders of dingoes, but it was
unknown to what extent that selection by breeders may have
influenced the genotypes of captive dingoes, or indeed if
hybrids existed in the pedigrees of the captive animals (Wilton,
Steward & Zafiris, 1999). In summary, current methods to
classify dingoes, feral dogs and dingo-dog hybrids based on
morphology, pelage and genetics appear to have poor dis-
criminatory abilities because natural variation within dingoes
is poorly understood; further, it is unknown if hybridization
may have altered the genome and phenotypes of the 20th and
21st century reference specimens.
A better description of the dingo, based on specimens that
are unlikely to have been influenced by hybridization, is
required to provide a benchmark against which to assess the
identities of dingoes in Australia. Such a description would
assist conservation and wildlife managers to classify dingoes
and to understand how the morphology of contemporary wild
Canis differs from pre-European dingoes. The purpose of this
paper is to provide that description.
Materials and methods
Specimens
Because Australia was colonized by Europeans in 1788 and
was only sparsely inhabited by European settlers prior to 1900
CE (Common Era) (Powell, 1991), we assumed that dingoes
collected prior to this date would be less likely to have been
influenced by hybridization with domestic dogs. We searched
the collections of museums held in Australia, Europe and the
US to locate dingo specimens that were known to or likely to
pre-date 1900 CE. The sample of 69 dingo skull specimens and
six skin specimens we subsequently located included speci-
mens taken by collectors in the 19th century and specimens
collected from archaeological and paleontological deposits
where museum data indicated that they pre-dated 1900 (Sup-
porting Information Table S1).We used radiocarbon (C14)
dating to determine if specimens from cave deposits that
lacked data on their context pre-dated 1900 (Supporting
Information Table S2). Radiocarbon dating for specimens
from the Western Australian Museum Palaeontology collec-
tions, 76.9.385, 76.9.384, 65.12.104, B3227b, B3227a, was
Figure 1 ‘Dog of New South Wales’ from Mazell & Phillip (1789).
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completed at Beta Analytic Radiocarbon Dating Laboratory,
Miami, Florida.
The selection of domestic dog C. familiaris specimens we
examined included breeds of similar size to dingoes which are,
or have frequently been used as stock-working dogs and
hunting dogs in Australia and thus could reasonably be
expected to have interbred with dingoes. These breeds
included Australian cattle dogs, kelpies, collies and grey-
hounds, and included specimens used by Newsome et al.
(1980).
Measurements
We took skull measurements with digital callipers (to the
nearest 0.01 mm) based on measurements given in Corbett
(1995), Macintosh (1975) and Von Den Driesch (1976)
(Table 1, Fig. 2). Additional measurements of Indian wolves
were obtained from Gollan (1982). Measurements for total
dingo series are given in Table 2.
Pelage coloration was recorded both from skins collected in
the 19th century which showed little discoloration from pres-
ervation or age, and from 18th century artists’ representations
of dingoes and early explorers and colonists’ reports of dingo
coloration. We based the coloration and markings criteria on
Elledge et al. (2008).
Diagnosis of dingoes from dogs
We first used stepwise discriminant function analysis to iden-
tify suitable measurements for the separation of dingoes from
dogs, producing a subset of 12 measurements for further
analysis. We then used a principal component analysis of
variables, standardized by size by dividing each measurement
by the geometric mean of all the measurements of that speci-
men (Mosimann, 1970), to investigate separation between
dogs and dingoes. We used canonical variates analysis to
quantify the separation of dingoes from dogs. We then com-
pared each individual dingo measurement to those of dogs
Table 1 Descriptions of measurements
Description Number
Length of auditory bulla (measured from where it abuts the paraocciptal process to the internal carotid foramen, excluding any
projection on the foramen)
1
Maximum maxillary width (measured at the junction of p4and Mlteeth) 2
Mid-crown width of P4tooth (measured through the highest cusp in a lateral direction) 3a
Greatest length of P43b
Greatest breadth of P43c
Basal crown length of C1(measured along tooth row) 4
Opisthion to inion (measured from a central inion point and not including the notch in the opisthion if present) 5
Width of both nasal bones (measured at premaxilla-maxilla suture) 6
Cranial height (measured from the upper notch of the external auditory meatus to the bregma, including the sagittal crest) 7
Distance between the posterior alveolar rims of Cl–P48
Height of the foramen magnum 9
Greatest breadth of the foramen magnum 10
Skull length (inion to prosthion which is the most anterior point on inter premaxillary suture) 11
Skull height (basioccipital to highest point of the sagittal crest) 12
Maximum post-orbital width (ectorbital to ectorbital) 13
Zygomatic breadth (zygion to zygion) 14
Viscerocranium length (nasion to prosthion) 15
Greatest neurocranium breadth =greatest breadth of the braincase (euryon to euryon) 16
Greatest mastoid breadth =greatest breadth of the occipital triangle (otion to otion) 17
Length of the cheektooth row (measured along the alveoli on the buccal side) 18
Condylobasal length: arboreal border of the occipital condyles to prosthion 19
Greatest length of Ml20a
Greatest breadth of Ml20b
Maxillar distance between P1and P1taken from the posterior alveolar rim 25
Dorsal width of incisive bone taken from mid-alveolar rims of canines 26
Greatest width of zygomatic arch taken from the base of suture between the zygomatic bone and zygomatic arch 27
Total length of palate 28
Premaxillary width between C1and C1taken from the alveolar rim across the base of the incisive bone 29
Basal length of the first incisor (measured along tooth row) 31
Premaxillary width taken between the two largest incisors taken from the alveolar rim across the base of the incisive bone 32
Length of parietal bone taken from central inion point on the sagittal crest to the bregma 34
Greatest width of auditory bulla 39
Maxillary distance between P4and P4taken from the anterior end of P4along the alveolar rim 41
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using analysis of covariance, with skull length as the covariate.
To enable easier diagnosis, and allowing for size, we plotted
each measurement against the total skull length.
Diagnosis
Differences from the wolf Canis lupus
The dingo differs from the wolf C. lupus, including the smaller
Indian wolf C. lupus pallipes, in being smaller in size in all
measurements (mean wolf condylobasal length =207.10 ±
2.10 s.e., mean pre-1900 CE dingo condylobasal length =
176.89 ±1.39; t90 =12.10, P<0.001). Dingoes also have more
variable pelage coloration, such as black and tan variants,
which are not found in wolves. Corbett (1995) shows separa-
tion of wolf skulls from dingo skulls using canonical variates
analysis, but does not give any scores, and included the larger
northern European and American wolves rather than the
Asian wolves from which dingoes were thought to be derived
(Oskarsson et al., 2011).
Differences from domesticated dogs C.
familiaris
There is some separation between dingoes and domesticated
dogs along PC2 in the size-adjusted principal component
analysis (Fig. 3), which accounts for 63.1% of the total vari-
ance (Table 3). This is mainly composed of a contrast between
Figure 2 Skull measurements used on dingo series.
Table 2 Measurements of pre-1900 CE dingoes
Measurement NMinimum Maximum Mean SD
2 Maximum maxillary width 69 52.10 70.80 59.03 3.69
25 Maxillary distance 47 26.08 37.77 31.70 2.67
29 Premaxillary width 45 17.22 27.23 21.42 2.01
19 Condylobasal length 63 147.25 199.80 176.89 11.06
8 Distance C1-P466 43.46 60.08 52.50 3.69
18 Length of the cheektooth row 61 54.62 72.31 64.24 3.57
4 Basal crown length of C151 8.56 12.19 10.12 0.94
31 Basal length of first incisor 33 4.45 7.94 6.13 0.69
28 Palate length 48 73.97 100.00 87.62 5.28
5 Opisthion to inion 65 37.90 55.33 47.42 3.74
15 Viscerocranium length 60 72.55 104.07 90.78 6.94
13 Maximum post-orbital width 54 40.58 63.51 51.51 4.51
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maximum post-orbital width and opisthion to inion length
with crown length of the first incisor and viscerocranium
length (Table 3). Canonical variates analysis did show some
separation for the non-size-adjusted measurements for domes-
ticated dogs and dingoes (Fig. 4), with differences largely
resulting from a contrast of viscerocranium length and
maximum maxillary width with condylobasal length, length of
cheektooth row and plate length (Table 4).
The dingo differs from the domestic dog C. familiaris and
its hybrids by restriction of pelage colours to combinations of
yellow, black and white, and in skull measurements including
relatively larger palatal width (Fig. 5a,c,g,j, Table 5), rela-
tively longer rostrum (Fig. 5e,f,i,k, Table 5), relatively shorter
skull height (Fig. 5b,d, Table 5) and relatively wider top ridge
of skull (Fig. 5h, Table 5).
Note that owing to the enormous variation in dog pheno-
types, dog breeds used in the analysis were restricted to those
of similar size and structure to dingoes.
Differences from other ‘dingo forms’
Note that the following canids are considered by some authors
as actual dingoes with some geographical variation (Corbett,
1985, 1995). Others recognized them as separate forms
(Gollan, 1982).
Figure 3 Plot of factor scores on first two axes of size-adjusted prin-
cipal component scores on pre-1900 CE dingoes and domestic dogs.
Table 3 Principal component loadings for size-standardized
measurements for dingoes and dogs
Component
123
2 Maximum maxillary width −0.841 −0.157 0.166
25 Maxillary distance −0.784 0.214 0.441
29 Premaxillary width −0.798 0.171 0.428
19 Condylobasal length 0.815 0.254 0.146
8 Distance C1-P40.836 −0.283 0.216
18 Length of the cheektooth row 0.783 −0.182 0.031
4 Basal crown length of C1−0.198 0.350 −0.681
31 Basal length of first incisor −0.142 0.712 −0.225
28 Palate length 0.744 0.367 0.297
5 Opisthion to inion 0.372 −0.495 0.160
15 Viscerocranium length 0.594 0.563 0.043
13 Maximum post-orbital width −0.109 −0.837 −0.278
Eigenvalue 5.053 2.303 1.182
% Variance 42.106 19.196 9.846
Figure 4 Canonical variate scores separating pre-1900 CE dingoes
from domestic dogs.
Table 4 Standardized canonical coefficients for measurements
separating pre- 1900 CE dingoes from domestic dogs
Function 1
2 Maximum maxillary width 0.420
25 Maxillary distance −0.097
29 Premaxillary width −0.018
19 Condylobasal length −0.823
8 Distance C1-P40.214
18 Length of the cheektooth row −0.802
4 Basal crown length of C10.018
31 Basal length of first incisor 0.379
28 Palate length −0.760
5 Opisthion to inion −0.631
15 Viscerocranium length 2.558
13 Maximum post-orbital width −0.085
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Figure 5 Bivariate plots comparing pre-1900 CE dingoes with domestic dogs with maximum skull length as Xaxis (a) maximum maxillary width, (b)
height of back of skull to inion on base of skull, (c) distance between the posterior alveolar rims, (d) basal crown length C1, (e) length of the cheektooth
row, (f) maxillar distance, (g) palate length, (h) width of top ridge, (i) length of nasal area, (j) premaxillary length, (k) distance C1–P4, (l) crown length
of first incisor.
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Figure 5 Continued.
M. S. Crowther et al. The Australian dingo
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1 Different from the New Guinea singing dog Canis
hallstromi by its greater height at the withers (Koler-Matznick
et al., 2003). It resembles the New Guinea singing dog in most
other morphological characteristics (Koler-Matznick et al.,
2003).
2 Different from Thai pariah dogs, as defined by Corbett
(1985), by being larger in cranial (total skull length of
pre-20th century dingoes 189.0 mm ±1.8; Thai pariah dog
male =179.5 mm ±3.1, female =173.2 mm ±3.6) and exter-
nal measurements (Corbett, 1985).
Description
Dingoes are dog-like and possess a fairly broad head, tapered
muzzle, erect ears and a bushy tail (Kerr, 1792; Fig. 6). Rela-
tive to similar-sized domestic dogs, dingoes have longer and
more slender muzzles. The 19th century dingoes we examined,
like wolves but unlike many dogs, do not possess dewclaws on
the hind legs (Ciucci et al., 2003). Dingoes can have five basic
pelage colours: yellow, brown, ginger/red, black and white
(Cairns, Wilton & Ballard, 2011). These colours occur in
various combinations and 19th century skin specimens
included animals that are entirely white (Fig. 6), entirely
yellow/brown (Fig. 6), entirely black, yellow with white
patches (Fig. 6), particularly at the tip of the tail and ankles
(Fig. 6), and yellow with black fur along the dorsal parts of the
body (sable, Fig. 6). The original specimen of C. dingo (Fig. 1)
illustrated in Mazell & Phillip (1789) was uniformly brown on
its dorsal surface, with the face, underparts and feet being
white (Kerr, 1792). Other pre-1800 paintings included colours
such as dark brown, reddish brown, and sandy with sabling
(Supporting Information Figure S1).
The specimen of C. macdonnellensis (Matschie, 1915) ZMB
22418 at the Museum für Naturkunde, Berlin, and the speci-
men of C. familiaris australasiae (Desmarest, 1820) at the
Muséum National d’Histoire Naturelle, Paris, were both pre-
dominantly yellow with some dark fur along the dorsum
(sabling).
Historical records describing dingo colours are scant, and
mostly not detailed (Elledge et al., 2006). The earliest report of
a dingo from 1697 is of a ‘yellow-dog’ near Jurien, Western
Australia (Abbott, 2008). Collins (1798) reported dingoes in
the Sydney region as ‘two colours, the one red with some white
about it, and the other quite black’. Explorer Mitchell (1839)
reported a ‘small black native dog’ in northern central New
South Wales in 1832. Historical descriptions of dingoes from
Western Australia during the period 1826–1890, compiled by
Abbott (2008), include red, yellow, black, black and white,
white, tan and tawny animals.
Mitochondrial and Y-chromosome DNA
Mitochondrial variation at the control region is posited to be
low in dingoes, with over 50% of animals sampled in previous
studies having a control region haplotype, A29, with all other
samples only differing by one base pair (Savolainen et al.,
2004; Oskarsson et al., 2011). This haplotype was shared with
dogs from East Asia, South-East Asian islands and Arctic
America (Savolainen et al., 2004). Similarly, only two
Y-chromosome haplotypes (H3 and H60) were found in
dingoes, the first shared with south-east Asian dogs and the
second derived from Taiwanese haplotypes, shared only with
the New Guinea singing dog (Ardalan et al., 2012). More
recently, dingoes have been found to exhibit a unique chro-
mosome haplogroup characterized by one single-nucleotide
polymorphism and 14 single tandem repeats (Sacks et al.,
2013).
Discussion
We have provided a morphological description of the dingo
based on specimens and information that are unlikely to have
been influenced by hybridization with domestic dogs. By pro-
viding a description for the dingo, our study provides a bench-
mark against which the identities of canids can be assessed.
Using our description, it is now possible to classify canids in
Australia as dingo-like based on morphological grounds.
Diagnosing the dingo
Diagnosing what constitutes a dingo remains difficult due to
the overlap in morphological characters with domestic dogs,
localized adaptations in dingoes and morphological variation
through time (Radford et al., 2012). Identification of diagnos-
tic morphological characters is also difficult, especially when
there is more variation within the domestic dogs in shape and
size than in the whole Canidae (Drake & Klingenberg, 2010).
Our morphological analyses showed that there is considerable
overlap between domestic dogs and dingoes for most morpho-
logical characters. This was particularly the case for some
Australian breeds, such as the Australian cattle dog, which are
thought to have dingo ancestry (Arnstein, Cohen & Meyer,
1964). A similar degree of overlap in shape exists between
North American wolves and closely related husky dogs
(Clutton-Brock, Kitchener & Lynch, 1994).
Consistent with previous studies, a broad cranium, widen-
ing of the palate and shortening of the rostrum were charac-
teristics separating domestic dogs from dingoes (Newsome
et al., 1980; Newsome & Corbett, 1982). Previous studies have
Table 5 ANCOVA values comparing cranial measurements on pre-1900
CE dingoes to domestic dogs with skull length (19) as covariate
Measurement Fd.f. P
2 Maximum maxillary width 5.20 1,104 0.025
25 Maxillary distance 25.71 1,80 0.052
29 Premaxillary width 5.15 1,80 0.025
8 Distance C1-P40.359 1,101 0.550
18 Length of the cheektooth row 5.67 1,96 0.019
4 Basal crown length of C10.05 1,82 0.824
31 Basal length of first incisor 11.34 1,64 0.001
28 Palate length 2.84 1,82 0.09
5 Opisthion to inion 1.81 1,100 0.182
15 Viscerocranium length 12.85 1,100 0.001
13 Maximum post-orbital width 2.92 1,93 0.091
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regarded widening of the palate and shortening of the rostrum
as indicators of domestication in dogs (Clutton-Brock, 2012).
The 19th century skin specimens we examined possessed erect
ears and a bushy tail. These characteristics were considered to
be typical traits of dingoes in the original description given by
Kerr (1792) and also in subsequent studies (Corbett, 1995;
Elledge et al., 2008). Pre-20th century dingoes lacked dew-
claws on the hind legs (Clutton-Brock, Corbet & Hills, 1976;
Ciucci et al., 2003).
The range of coat colours that can occur in dingoes is a
controversial subject, with some authors only accepting black,
and black and tan dingoes (Macintosh, 1975; Newsome &
Corbett, 1985; Jones, 2009), while others only accepting
yellow or light brown (ginger) and rejecting animals with dark
dorsal fur (sable) (Elledge et al., 2008). The small sample of
19th century dingo skins and 18th century illustrations of
dingoes we examined shows that there was considerable vari-
ability in the colour of dingoes, and that their coloration was
not restricted to just yellow and white animals, but also
included various combinations of yellow, white, brown and
black. The range and combinations of coat colours in these
skins and illustrations were consistent with historical accounts
Figure 6 Colour variation in museum specimens of dingoes: (a) ginger collected between 1801 and 1803 (MNHN_2 M 2010.671), (b) sable collected
1820 (MNHN_2 M 2010_672), (c) black collected in the early 19th century (BMNH 50.11.30.22), (d) black collected in 1856 (C1829), (e) sable
collected in 1842 (BMNH 1842.9.19.1), (f) white collected in the late 19th century (BMNH 1939.1697). Specimens (a) and (b) are from the Muséum
National d’Histoire Naturelle, Paris; (c), (e) and (f) are from the Natural History Museum, London; and (d) is from the Museum Victoria.
M. S. Crowther et al. The Australian dingo
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from the 19th century and observations of dingoes made by
Newsome & Corbett (1985). Markings such as white spots
restricted to feet, chest spot, neck flash, underbelly and tail tip,
as used by the Australian National Kennel Council in
the dingo breed standard (http://www.ankc.org.au/Breed_
Details.aspx?bid=103), are not recorded in most early
accounts, and are not present in all pre-1900 CE skins or
illustrations.
The presence of individuals with sable pelage (dark dorsal
coloration and lighter lateral coloration: Fig. 6b,d) in the
sample of 19th century skins suggests that this coloration is
not necessarily indicative of hybridization as has been sug-
gested by previous authors (Corbett, 1995; Elledge et al.,
2008). The sample of skins and illustrations we examined did
not include animals with brindle coloration. Brindle, dingo-
like dogs appear in the historical record from the 1890s, and
could plausibly be the result of hybridization, particularly as it
is a colour pattern found in greyhounds, which were brought
into Australia in 1788 and are not found in most older dog
breeds (Cairns et al., 2011). However, the small sample size of
specimens we examined does not allow inferences to be made
as to whether brindle individuals are dingo-dog hybrids or
dingoes.
C. lupus dingo,C. familiaris dingo
or C. dingo?
There has long been a confusion regarding the identities and
classification of wild mammal species and their descendent
domestic forms (Gentry, Clutton-Brock & Groves, 1996).
Many authors classify domesticates as subspecies of the
species from which they are thought to be descended (Wilson
& Reeder, 2005). Following Corbett (1995), most recent
authors quote the dingo as C. lupus dingo on the assumption
that they, along with domestic dogs, were descended from a
common ancestor, the grey wolf C. lupus. However, recent
research has suggested that C. lupus is a species complex,
which contains distinct clades and cryptic species (Aggarwal
et al., 2007; vonHoldt et al., 2011), and that C. familiaris and
C. dingo do not fall within any modern wolf clade (Freedman
et al., 2014). In addition, as domesticated forms do not fall
into the definition of subspecies, the ICZN has recommended
retaining the different specific names for wild and domesti-
cated animals and naming wild ancestors of domesticates
using the first available specific name based on a wild popu-
lation (ICZN, 2003). Hence, we argue that because the ances-
try of the dogs and dingoes is unknown, and because the dingo
was first described as a distinctive wild form and differs from
wolves, New Guinea singing dogs and domestic dogs in many
behavioural, morphological and molecular characteristics
(Macintosh, 1975; Corbett, 1995; Wilton et al., 1999), and
they are effectively reproductively isolated in undisturbed
natural environments and thus like C. hallstromi can be con-
sidered a distinct taxon (Koler-Matznick et al., 2003).
Furthermore, because the dingo was first described as C. dingo
Meyer 1793, and this decision was later upheld by ICZN
(1957), we propose that C. dingo is the correct binomial.
Conservation and management
Our study reveals that the pelage criteria used in previous
studies to diagnose dingoes (Newsome & Corbett, 1985;
Elledge et al., 2008) do not encompass the morphological vari-
ation present in pre-20th century specimens. Many managers
currently cull animals they believe to be hybrids based on
pelage coloration. In particular, animals with sable pelage are
frequently culled because they do not conform with previous
criteria used to define dingoes (M. Letnic, pers. obs.). Our
findings suggest that such culling may be unwarranted because
animals with this coloration appear in the illustrations and
skin specimens from 18th and 19th centuries (Fig. 6). Indeed,
there is a risk that the use of pelage to diagnose dingoes may
result in humans selecting for yellow dingoes because this
common colour morph of dingoes is widely perceived as being
the colour of ‘pure’ dingoes (Elledge et al., 2006). The next
step for the conservation and integrity of dingoes is to define
characters to separate dingoes from hybrids, allowing for
natural selection and recognizing the variation naturally
present in dingoes.
Acknowledgements
We thank the many staff from museums for providing access
to their collections. Funding was provided by the Asia Pacific
Science Foundation. Kylie Cairns and Chris Dickman com-
mented on a draft. Anna Feit translated German texts.
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Supporting information
Additional Supporting Information may be found in the
online version of this article at the publisher’s web-site:
Figure S1. Pre-1800 paintings of Australian dingoes. (a) A
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Table S1. List of pre-1900 CE dingo specimens used in
analyses.
Table S2. Dates of previously undated dingo cave specimens.
The Australian dingo M. S. Crowther et al.
12 Journal of Zoology •• (2014) ••–•• © 2014 The Zoological Society of London