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The denning behaviour of dingoes (Canis dingo) living
in a human-modified environment
Bradley Smith
A,C
and Anne-Louise Vague
B
A
School of Human Health and Social Sciences, CQUniversity Australia, Building 32/2.27,
Bruce Highway, North Rockhampton, Qld 4702, Australia.
B
Newcrest Mining Limited, Telfer Environment, 234 Railway Parade, West Leederville, WA 6007, Australia.
C
Corresponding author. Email: b.p.smith@cqu.edu.au
Abstract. Little is known about reproduction and den site selection by free-ranging dingoes. We present observations
of den sites used by dingoes inhabiting a large-scale mining operation located in the Great Sandy Desert, Western Australia.
We observed 24 dens concentrated within a 1-km radius. Den sites were generally situated in elevated positions overlooking
the surrounding area, were a short distance from food and water resources, required vegetation (particularly spinifex grass)
to provide a firm foundation and stable ceiling in the soft sand, and had single den openings that faced away from the rising
and daytime sun. Distance to human structures or activity did not appear to influence site selection. Four of the dens were
active, containing a total of 37 pups aged between two and four weeks of age. One den contained 18 pups of different ages,
indicating that communal denning was also occurring. The high number of breeding females within close proximity
suggests that multiple family groups are able to share resources and live in close proximity. Our findings highlight the
importance of human-modified areas and abundance of resources in the reproduction and breeding site selection of dingoes.
Additional keywords: breeding, Canis lupus, den site, dingo, reproduction, wild dog.
Received 29 February 2016, accepted 18 August 2016, published online 27 September 2016
Introduction
For wild canids, selection of den sites, and the activities that occur
at den sites is directly related to the reproductive success of the
pack (Fuller et al. 2003; Benson et al. 2015). For grey wolf pups
(Canis lupus), where mortality is highest in the first six months of
life (Harrington and Mech 1982; Benson et al. 2013), adaptive
selection of den sites should maximise the ability of the pack to
efficiently provision the breeding female and her pups, as well as
provide safety from predators and other threats (Benson et al.
2015). Knowledge of den selection and characteristics can benefit
conservation and management by providing insight into the
relationship between environmental conditions and population
dynamics. Although our understanding of grey wolf reproductive
and denning behaviours is quite comprehensive (Mech 1970),
reports directly relating to such denning behaviours in the dingo
(Canis dingo), a wild canid endemic to Australia (Smith 2015)
are scant, with the exception of ad hoc observations by
Thomson (1992a), Corbett and Newsome (1975), Harden (1981),
and Breckwoldt (1988).
Like grey wolves, male and female dingoes can exhibit long-
term pair bonding, maintain and defend a shared territory,
cooperatively care for young, and reproduce seasonally (Corbett
and Newsome 1975; Thomson 1992a, 1992b, 1992c; Corbett
2001; Lord et al. 2013; Smith 2015). Dingo pups are generally
born in dens in the winter months (June to August), which
coincides with periods that are optimal for rearing young (Smith
2015). The average litter size is 4–5 pups, with a possible range
of 1–10 (Corbett and Newsome 1975; Jones and Stevens 1988;
Catling et al. 1992; Corbett 2001; Allen et al. 2015; Smith 2015).
Most dingoes do not become sexually mature until their
second year, although in captivity, and/or areas where resources
are abundant, it can occur earlier (Catling 1979; Catling et al.
1992; Corbett 2001; Smith 2015). It is generally believed that
only one dingo bitch has pups per territory (Smith 2015);
however, this is not always the case. On Fraser Island, for
instance, multiple litters and multiple lactating females have been
observed in the same pack area or territory (Allen et al. 2015).
In many respects, dingoes’choice of den site mirrors that of
wolves (Mech 1970; Kowalewski 2009). Dingo dens have been
found in hollow logs, enlarged rabbit warrens and rock shelters,
under large tussocks of spinifex and at the bases of large trees
(Thomson 1992a;Corbett 2001). The most comprehensive
study relating to dingo den selection was conducted by Thomson
(1992a), who reported observations of 21 natal dens and pup
rearing in wild dingos around Fortescue River, Western Australia,
between 1975 and 1984. In this region, dingoes appeared
selective in their choice of sites, showing preference for
elevated sites (that often commanded extensive views of lower
approaches), proximity to water (within 2.5 km, but most within
0.5 km), and den openings oriented away from the north to limit
Journal compilation ÓAustralian Mammal Society 2016 www.publish.csiro.au/journals/am
CSIRO PUBLISHING
Australian Mammalogy
http://dx.doi.org/10.1071/AM16027
the excessive heat from the sun. Dens were found in everything
from rock piles, ledges and outcrops, shallow hollows underneath
a large tussock of spinifex, hollow logs, and an enlarged burrow
of the large sand monitor (Varanus gouldii). In hilly areas,
dingoes appeared to prefer to create tunnels that had multiple
smaller cavities, or tunnels leading from the main chamber that
extended for several metres. The main chambers generally
opened directly to the outside, with some having multiple
entrances. Although the size of the entrances differed, most were
large enough to allow a human to crawl inside.
The availability of resources, such as food and water, are key
drivers of dingo distribution and behaviour (Smith 2015).
Anthropogenic activity such as tourism (e.g. at locations such as
Fraser Island, and Uluru: Allen et al. 2015; Behrendorff et al.
2016), periurban areas (Allen et al. 2016), and mining operations
(Newsome et al. 2013a) that occur within natural dingo home
ranges often provide supplementary resources that can influence
dingo distribution and behaviour, and lead to negative
human–dingo interactions or conflict (Lawrance and
Higginbottom 2003; Appleby 2015). Newsome et al. (2013a),
for example, discovered that average group sizes, genetic
diversity, and diet in a population of dingoes living around two
mines in the Tanami Desert, Northern Territory, were greater in
areas with abundant focal food sources around the mine sites,
compared with those in more distant areas. This provided support
for the notion that resource richness favours larger group size,
consistent with the Resource Dispersion Hypothesis (Macdonald
1983). Although the findings from the report by Newsome
et al. (2013a, 2013b, 2014) reinforce the notion that access
to anthropogenic resources alters the social structure and
movements, the impacts of extreme resource concentration on
dingo reproductive behaviour such as the selection and density
of natal dens remains uncertain.
In a first step to rectifying this, we provide an account of the
denning behaviour of dingoes living in a human-modified
environment (a mining operation). Such information is pertinent
to gaining a greater understanding of the species, and to
monitoring and managing dingo populations.
Materials and methods
Study area
Newcrest Mining Limited’s Telfer Copper and Gold Mine
(Telfer) is located in the Great Sandy Desert, Western Australia
(21.71S, 122.23E). The annual mean temperature is 34.1C
(highest monthly mean of 37.4C, and lowest monthly mean of
30.1C). In June and July (winter/dry season), during the height
of the whelping season, the mean temperature is 25.3C
(highest monthly mean of 27.9C, lowest monthly mean of
23.4C). During December and January (summer/wet season),
the mean temperature is 40.4C (highest monthly mean of
43.5C, lowest monthly mean of 36.4C). Average annual
rainfall is 30.4 mm, with the average during June and July
being 13.2 mm, and the average during December and January
being 49.9 mm (Telfer Aero, site 013030, based on data from
1974–2015: Australian Bureau of Meteorology 2016).
Telfer Mine comprises open pit and underground operations
and there are ~800 employees and contractors that operate on a
‘fly-in-fly-out’basis (typical shift rotation involves eight days
on site, six days off site). The site operates 24 h per day, seven days
per week (with day- and night-shift rosters). Mine infrastructure
includes water storage ponds, water pipelines, sewerage treatment
plant, power station, administration buildings and work sheds
throughout the site, underground mine, two open pits, large piles
of waste rock, topsoil storage areas, and a landfill (where all
waste products are disposed, including all food waste). Facilities
at the village include an airfield, administration buildings, dining
hall and kitchen, BBQ cooking facilities, accommodation
rooms, and sporting facilities (swimming pool, golf range, tennis
courts). The mine site area covers 4910 ha. The closest human
settlement to Telfer is an Indigenous community located
~100 km away.
Dingo population
The main habitats in this region of the Great Sandy Desert are
sand plains, stony hills, linear sand dunes and swales, and isolated
clay pans. The region includes a wide variety of vertebrate fauna,
the following having been verified through analysis of dingo scats
collected on site (Smith, unpubl. data): reptiles (e.g. Scincidae,
Varanidae, Elapidae), birds (e.g. Dromaius novaehollandiae);
feral mammals such as camel (Camelus dromedarius), cat (Felis
catus), fox (Vulpes vulpes), and small marsupials and rodents (e.g.
Pseudomys hermannsburgensis,Zyzomys argurus,Dasykaluta
rosamondae). A comprehensive and current estimate of the
dingo population has yet to be conducted, although we estimate
that the population during the 2015 whelping season was ~120
individuals across the mine footprint (excluding the pups born in
2015). Population estimates were determined through observing
and subsequent counting of dingoes by the authors at various
locations around the mine site over a period of 18 days (nine days
in July and nine days in December 2015), and based on the
outcomes of a cull conducted in February 2016. Genetic testing
of a random sample of the resident dingo population (n= 44,
conducted by Zoological Genetics in 2015) indicates that the
population has a high level of genetic purity (average = 92.6%,
range = 82–99%). Historically, dingo numbers had been managed
by culling onsite when there were families and domestic pets in
the town. Since the Telfer expansion in 2004 the focus has
been on minimising interaction between dingoes and humans,
minimising dingo access to food (e.g. education of staff, fencing
eating areas, locking bins), and ad hoc culling programs.
Den sites
Dens (active and inactive) were visited over a 5-day period in
July 2015. Searches on foot were conducted in and around the
immediate vicinity of the mine site to identify areas of high dingo
activity. A total of 19 search areas (of various sizes) were selected
(Fig. 1) according to recent and past observation of dingo activity
identified by mine staff. High dingo activity was determined
according to the presence of footprints/tracks in the sand,
trampled vegetation, and scats. Six areas showed no activity, four
showed limited or old activity, six showed high activity but were
considered unlikely to contain a den site (e.g. transit areas), and
three areas showed high levels of recent activity and were
considered likely to contain a den site A detailed transect was
conducted of these areas, and were subsequently labelled Area A
(10.98 ha), Area B (22.06 ha) and Area C (13.91 ha) (Fig. 1).
BAustralian Mammalogy B. Smith and A.-L. Vague
The outside of the dens were photographed before the area
was disturbed by the researchers, and the location recorded using
a hand-held GPS device (Garmin eTrex H, Garmin International
Inc., KS, USA). A bearing (Garmin eTrex H) from the main
entrance was taken, as well as photographs of inside the den. The
width and depth of main chamber was measured using a 5-m
retractable steel tape measure. Lastly, we noted the position of
the den opening according to its position in the landscape (e.g.
ground level, or position on a mound), species of vegetation
surrounding the den, and whether any canopy was present.
The distances from the den to fresh water, roads, buildings, and
rubbish tip were measured using ArcGIS software after den
GPS positions were entered into the software. If pups were
present, they were counted during examination of the den, or by
observing the pups outside the den. This method enabled
complete counts to be made. Age was estimated according
to physical characteristics (size and open/closed eyes) and
behavioural development (the ability to walk, and the dexterity of
(a)
(b)
Fig. 1. (a) An aerial view of the mine site. Areas highlighted green indicate no dingo activity, blue indicates
old/limited dingo activity, yellow indicates high activity but unlikely to contain a den site, and red indicates
high levels of activity and considered likely to contain a den site. (b) Location of dingo natal dens in relation to
roads, water source, landfill, and mining operations. Area A, topsoil storage area (Dens 1–16); Area B, topsoil
storage area and natural bushland (Dens 17–23); and Area C, the landfill (Den 24). Green dots indicate active
dens, and red dots inactive dens.
Dingo den site selection Australian Mammalogy C
movement) using comparative data from studies of dingoes
(Smith 2015).
No more than 10 min was spent at each den site, and care was
taken to minimise the disturbance to the den and to the pups (i.e.
they were not handled). No adult dingoes were present, but
two were nearby and fled on approach. Despite this, out of the
four active dingo den sites visited, only two litters remained in the
den on the following days. Such a response by dingoes was
also observed by Thomson (1992a). It is therefore important
when studying den sites to realise that, like other canids (e.g.
grey wolves: Mech 1970; Kowalewski 2009; Person and Russell
2009), dingoes are sensitive to human encroachment.
Results
Den characteristics
We located 24 den sites across three sites within the mine (Fig. 1).
Most of the dens were located in topsoil storage areas (79%,
n= 19). Prior to any land clearing (e.g. mining, roads and waste
dumps), the topsoil of the disturbance area is collected (by
stripping with a loader) to a depth of 250 mm. This topsoil is
relocated to the closest topsoil storage area for future progressive
rehabilitation works. Topsoil is ‘paddock dumped’into mounds
that do not exceed 3 m in height. These mounds provided the
ideal location for dingo natal dens. The remainder were located
in natural/undisturbed bushland (17%, n= 4), and one was
located in the landfill (4%, n= 1). All dens were located within a
1-km radius of each other, and different distances to the nearest
unsealed road (102 m 11 s.e.), human structure (343 m
27 s.e.), major water source (302 m 33 s.e.), and landfill
(731 m 38 s.e.).
Of the 24 dens located, two were located at ground level, two
at the top of natural mounds, one at the landfill, and 19 within
an artificial (topsoil) mound. The dens positioned in topsoil
mounds varied in their position. These included within the
bottom third of a mound (37%, n= 7), base of a mound (21%,
n= 4), top third (16%, n= 3), top of mound (16%, n= 3), and
half-way (10%, n= 2) (see Table 1). All dens were surrounded
by, or close to, native grasses (typically either soft spinifex
(Triodia pungens) or hard spinifex (Triodia basedowii)). The
roots of the grasses and/or trees provided a firm foundation
and stable ceiling in the sandy soil. The other common
vegetation included small (~1–2-m high) native trees from the
genus Acacia, which provided some coverage for four of the 23
natural dens (17%). The den in the landfill was also protected;
however, the coverage consisted of hard rubbish (e.g. large
wooden crates). Common to all active dens was evidence of
high dingo activity: for example, footprints that formed networks
of pathways, resting pads/beds, scats, and anthropogenic objects
(e.g. leather boots and gloves) that had evidence of chew marks.
Above or beside the den there was often an observation post
overlooking a large open area. Fig. 2 presents a typical example
of a den in this area.
The openings of the dens, on average, measured 456 mm
wide (37 s.e., and 421 mm in height (27 s.e.) and were
typically domed (Fig. 2). The average depth of the chamber
measured 1045 mm (70 s.e.). Dens had only one opening,
with the exception of Den 16, which had multiple entrances.
The den located in the landfill (Den 24) was positioned within
Table 1. Descriptions of dingo natal dens including the number of dens, the number of dens with a canopy, and the number of active dens grouped according to the position of den in
relation to the ground
Mean (s.e.m.) dimensions (cm) are provided for the height and width of the den openings, the depth of the main chamber, as well as the elevation (m). Distances (m) to the nearest road, human structure,
water source and landfill are also included (mean s.e.m)
Position of den NCanopy Active Den Dimensions Distances (m) to:
Height (cm) Width (cm) Depth (cm) Elevation (m) Road Human structure Water source Land fill
Ground 2 0 0 385.00 ± 5.00 505.00± 115.00 1020.00 ± 380.00 298.5 ± 2.50 45.50 ± 6.50 358.50 ± 109.50 354.50 ± 235.50 692.00 ± 88.00
Base of mound 4 0 1 488.75 ± 79.49 370.00 ± 111.43 1070.00 ± 261.85 295.75 ± 2.50 176.50 ± 6.60 293.25 ± 39.99 265.00 ± 32.88 796.75 ± 48.95
Bottom 1/3 of mound 7 1 0 450.00 ± 71.48 416.85 ± 62.27 962.85 ± 148.54 297.71 ± 1.81 117.14 ± 17.95 358.00 ± 45.15 288.57 ± 56.86 788.14 ± 50.00
Half way up mound 2 1 1 405.00 ± 95.00 530.00 ± 10.00 975.00 ± 75.00 263.50 ± 34.50 80.50 ± 13.50 383.50 ± 223.50 334.00 ± 134.00 706.50 ± 101.50
Top 1/3 of mound 4 1 1 382.50 ± 28.39 595.00 ± 113.83 1252.50 ± 104.67 297.75 ± 1.03 70.75 ± 19.12 333.25 ± 58.39 264.00 ± 104.63 770.00 ± 48.66
Top of mound 4 1 0 366.25 ± 29.25 410.00 ± 61.78 1002.50 ± 66.76 297.50 ± 2.78 89.50 ± 22.62 385.25 ± 83.99 352.25 ± 102.20 739.75 ± 52.65
Landfill 1 1 1 –– – –20.00 197.00 335.00 –
DAustralian Mammalogy B. Smith and A.-L. Vague
a complex network of large wooden crates, which prevented
accurate measurements.
Most dens, including active dens (those containing pups), had
openings that faced in a south-west direction (bearing 190–264)
(Fig. 3). This was away from the rising sun (bearing ~58).
Reproduction
Four of the dens located contained pups that were estimated to be
between two and four weeks of age. Thus, mating likely occurred
in April, and whelping in June. Estimated mating date for Dens 4,
8 and 21 was 1 April 2015 (based on a whelping date of 3 June
2015), and estimated mating date for Den 24 was 15 May 2015
(based on a whelping date of 17 June 2015) (mating date was
calculated by subtracting 63 days, the average gestation time,
from the estimated whelping date). One of the dens contained 18
pups (see Fig. 4aand b) but, given the variation in pup size and
locomotion, it likely consisted of three different litters (aged two
to four weeks). Overall, this suggests that there were at least 4–6
breeding females, and 37 pups in close proximity during the
2015 breeding season. Although whelping occurs in a defined
period, it is highly likely that more litters were born after the
survey, and thus this estimate is a conservative one.
Discussion
This paper provides an observational account of the denning
behaviour and site selection of a population of free-ranging
dingoes living in an area where human-related sources of food
and water are abundant throughout the year. Uniquely, at this site,
a high concentration of dens and puppies were located within a
small area (within 1 km). However, the characteristics of the
dens appear similar to those reported for dingoes elsewhere
(Corbett and Newsome 1975; Harden 1981; Breckwoldt 1988;
Thomson 1992a; Corbett 2001), as well as for grey wolves (Mech
1970; Person and Russell 2009; Kowalewski 2009). Some
notable exceptions here include the lack of multiple entrances, the
lack of diversity in terms of den ‘types’, and the sheer number of
dens in close proximity. In the Great Sandy Desert, the range
of den types is somewhat limited. However, it appears that the
artificial creation of topsoil storage areas provides a suitable
material for den construction, with the position on the mound
being related to local conditions (such as sun, wind, and
vegetation). The lack of predators in the area may have reduced
the need for multiple den entrances.
It is not known why there was such a high number of den
sites in close proximity. Thomson (1992a) and Harden (1981)
suggest that dingoes move den sites to follow food sources.
However, at the mine, resources were available year-round.
High numbers of dens might indicate a high rotation of den sites
each year, as also reported by Harden (1981). Pups might be
regularly moved after birth, or perhaps a new den is preferred
each breeding season and den sites are not reused (as has been
observed in a grey wolf population in Finland: Kaartinen et al.
2010). It is not known whether the dens were being used outside
of the breeding season as a way for adults to seek protection
from the extreme heat of the day, and/or the rains in the wet
season.
Although wild canids tend to establish den sites away from
roads or areas of high human activity, they are resilient and can
persist in human-dominated landscapes (Person and Russell
2009; Ahmadi et al. 2014). Dingoes, for instance, are capable
of successfully living within periurban areas (Corbett 1998;
Allen et al. 2013), yet little is known about their denning
behaviour in such areas. In the current study, dens were located
close to man-made structures and activity. Human-disturbed
areas such as Fraser Island (Allen et al. 2015), and mining
operations in remote areas (Newsome et al. 2014), appear to
provide the ideal environment for dingo subsistence. At this
site, for example, mining operations provide constant access to
food (via the landfill and accommodation village), ample water
Fig. 2. A typical den in the topsoil storage area. Most dens had arched
openings, were positioned below vegetation (in this case spinifex) to provide a
firm foundation and stable ceiling in the sandy soil, and provided elevated
areas that acted as lookouts.
0°
2
N
E
S
W
15°
30°
45°
60°
75°
90°
105°
120°
135°
165°
180°
195°
210°
225°
270°
300°
315°
330°
345°
1
240°
255°
285°
150°
Fig. 3. The orientation of main entrances grouped according to direction.
The asterisk (*) indicates active natal dens (Den 4, 213; Den 8, 205; Den
21, 243). The length of each bar reflects a frequency of 1 (inner circle) or
2 (outer circle).
Dingo den site selection Australian Mammalogy E
(a)(b)
(c)(d)
(e)(f)
(g)(h)
Fig. 4. View of the outside and inside of each active den. (a,b) Den 4 containing seven pups aged 3–4 weeks, located at ground level;
(c,d) Den 8 containing 18 pups of various ages (2–4 weeks) (a small ledge outside the den opening provided a barrier preventing smaller
pups from straying from the den); (e,f) Den 21 containing six pups aged 3–4 weeks, located halfway up a topsoil mound and covered by
dead Acacia trees; (g,h) Den 24 containing six pups aged two weeks, located in the landfill underneath large wooden crates and boxes.
FAustralian Mammalogy B. Smith and A.-L. Vague
(available through dripping or leaking pipes, and water storage
ponds of various sizes), navigation channels (created through
the establishment of dirt roads) and shade (available throughout
the site beneath oversize truck tyres, pipes and buildings),
and allow the construction of dens (e.g. topsoil storage areas).
Despite the location of the den sites being close to high human
activity, they were rarely, if ever, disturbed by the miners. It
appears that dingoes have habituated to the presence of people
and vehicles, and to the constant noise and activity on site. Thus,
the human-disturbed landscape in this population can be seen
to be helping, not hindering, the survivability of dingoes.
As in free-ranging dingo populations observed in central
Australia (Green and Catling 1977) and Fraser Island (Allen
et al. 2015), we discovered several breeding females raising
litters within the same general area/territory. In addition, we
observed an incidence of communal denning. This raises
questions about the prominence of the role of infanticide by the
dominant female, as proposed by Corbett (1988), which has
been observed in only a single captive population (Corbett
1988). It is more than likely that the conditions of Corbett’s
study site (limited enclosure size, increased aggression, and
lack of opportunity for individuals to disperse) gave rise to this
abnormal behaviour, or at least the probability of it being
observed.
The findings reported here are observational only, and lack
denning and reproductive behaviour of free-ranging dingoes
from nearby unmodified sites. Nonetheless, our observations
highlight the significance of human-modified areas and
abundance of resources (human-related sources of food and
water) in the reproduction and breeding site selection of
dingoes. That is, dingo numbers are likely to swell, and multiple
family groups are able to live in close proximity around shared
resources. It is imperative that access to resources available to
dingoes (such as food in landfills and where humans frequent)
continues to be a management priority.
Acknowledgements
Funding for this study was provided by Telfer Mines Environmental
Department (Newcrest Mining Limited), and from an internal grant from
Central Queensland University. The data presented in this study were
collected by Telfer Environment staff as part of their ongoing monitoring
and management of the resident dingo. Permission to access, analyse and
publish the data was sought from, and subsequently approved by, the Central
Queensland University Animal Ethics Committee, Project No. A15/09-335.
The authors thank Chris Tiemann, Matthew Shaw and Claire Learey for
assistance during fieldwork, and Dr Damian Morrant, Robert Appleby,
Dr Ben Allen, Dr Ross Goldingay and an anonymous reviewer for valuable
comments on the manuscript.
References
Ahmadi, M., López-Bao, J. V., and Kaboli, M. (2014). Spatial heterogeneity
in human activities favors the persistence of wolves in agroecosystems.
PLoS One 9, e108080. doi:10.1371/journal.pone.0108080
Allen, B., Goullet, M., Allen, L., Lisle, A., and Leung, L. (2013). Dingoes
at the doorstep: preliminary data on the ecology of dingoes in urban
areas. Landscape and Urban Planning 119, 131–135. doi:10.1016/
j.landurbplan.2013.07.008
Allen, B., Higginbottom, K., Bracks, J., Davies, N., and Baxter, G. (2015).
Balancing dingo conservation with human safety on Fraser Island: the
numerical and demographic effects of humane destruction of dingoes.
Australasian Journal of Environmental Management 22, 197–215.
doi:10.1080/14486563.2014.999134
Allen, B., Carmelito, E., Amos, M., Goullet, M., Allen, L., Speed, J., Gentle,
M., and Leung, L. (2016). Diet of dingoes and other wild dogs in peri-
urban areas of north-eastern Australia. Scientific Reports 6, 23028.
doi:10.1038/srep23028
Appleby, R. (2015). Dingo–human conflict: attacks on humans. In ‘The
Dingo Debate: Origins, Behaviour and Conservation’. (Ed. B. Smith.)
pp. 131–158. (CSIRO Publishing: Melbourne.)
Australian Bureau of Meteorology (2016). Monthly climate statistics for
‘TELFER AERO’(013030). Available at: http://www.bom.gov.au/
climate/averages/tables/cw_013030_All.shtml [accessed 29 February
2016].
Behrendorff, L., Leung, L. K.-P., McKinnon, A., Hanger, J., Belonje, G.,
Tapply, J., Jones, D., and Allen, B. L. (2016). Insects for breakfast
and whales for dinner: the diet and body condition of dingoes on Fraser
Island (K’gari). Scientific Reports 6, 23469. doi:10.1038/srep23469
Benson, J. F., Mills, K. J., Loveless, K. M., and Patterson, B. R. (2013).
Genetic and environmental influences on pup mortality risk in a
Canis hybrid zone. Biological Conservation 166, 133–141. doi:10.1016/
j.biocon.2013.06.018
Benson, J. F., Mills, K. J., and Patterson, B. R. (2015). Resource selection
by wolves at dens and rendezvous sites in Algonquin Park, Canada.
Biological Conservation 182, 223–232. doi:10.1016/j.biocon.2014.
12.010
Breckwoldt, R. (1988). ‘A Very Elegant Animal: The Dingo.’(Angus and
Robertson: Sydney.)
Catling, P. (1979). Seasonal variation in plasma testosterone and the testis
in captive male dingoes, Canis familiaris dingo. Australian Journal of
Zoology 27, 939–944. doi:10.1071/ZO9790939
Catling, P., Corbett, L. K., and Newsome, A. (1992). Reproduction in
captive and wild dingoes (Canis familiaris dingo) in temperate and
arid environments of Australia. Wildlife Research 19, 195–209.
doi:10.1071/WR9920195
Corbett, L. K. (1988). Social dynamics of a captive dingo pack: population
regulation by dominant female infanticide. Ethology 78, 177–198.
doi:10.1111/j.1439-0310.1988.tb00229.x
Corbett, L. (1998). Management of dingoes on Fraser Island. CIC prepared
for Queensland Department of Environment by ERA Environmental
Services Pty Ltd.
Corbett, L. K. (2001). ‘The Dingo in Australia and Asia.’(JB Books:
Adelaide.)
Corbett, L. K., and Newsome, A. (1975). Dingo society and its maintenance:
a preliminary analysis. In ‘The Wild Canids: Their Systematics,
Behavioral Ecology and Evolution’. (Ed. M. W. Fox.) pp. 369–379.
(Van Nostrand Reinhold: New York.)
Fuller, T. K., Mech, L. D., and Cochrane, J. F. (2003). Wolf population
dynamics. In ‘Wolves: Ecology, Behavior, and Conservation’. (Eds
L. D. Mech and L. Boitani.) pp. 161–191. (University of Chicago Press:
Chicago, IL.)
Green, B., and Catling, P. (1977). The biology of the dingo. In ‘Australian
Animals and their Environment’. (Eds H. Messel and S. T. Butler.)
pp. 51–60. (Shakespeare Head Press: Sydney.)
Harden, R, (1981). A look at the dingo. Australian Natural History 20,
191–194.
Harrington, F. H., and Mech, L. D. (1982). Patterns of homesite attendance
in two Minnesota wolf packs. In ‘Wolves of the World: Perspectives
of Behavior, Ecology, and Conservation’. (Eds F. H. Harrington and
P. C. Paquet.) pp. 81–105. (Noyes Publications: Park Ridge.)
Dingo den site selection Australian Mammalogy G
Jones, E., and Stevens, P. (1988). Reproduction in wild canids, Canis
familiaris, in the eastern highlands of Victoria. Australian Wildlife
Research 15, 385–394. doi:10.1071/WR9880385
Kaartinen, S., Luoto, M., and Kojola, I. (2010). Selection of den sites by
wolves in boreal forests in Finland. Journal of Zoology 281,99–104.
doi:10.1111/j.1469-7998.2009.00678.x
Kowalewski, D. (2009). The anatomy of a wolf den site: a field report.
Electronic Green Journal 1(28).
Lawrance, K., and Higginbottom, K. (2003). Behavioural responses of
dingoes to tourists on Fraser Island. CRC for Sustainable Tourism.
Lord, K., Feinstein, M., Smith, B., and Coppinger, R. (2013). Variation
in reproductive traits of members of the genus Canis with special
attention to the domestic dog (Canis familiaris). Behavioural Processes
92, 131–142. doi:10.1016/j.beproc.2012.10.009
Macdonald, D. W. (1983). The ecology of carnivore social behaviour. Nature
301, 379–384. doi:10.1038/301379a0
Mech, L. D. (1970). ‘The Wolf: Ecology and Behaviour of an Endangered
Species.’(Natural History Press: New York.)
Newsome, T. M., Ballard, G. A., Dickman, C. R., Fleming, P. J., and van de
Ven, R. (2013a). Home range, activity and sociality of a top predator,
the dingo: a test of the Resource Dispersion Hypothesis. Ecography 36,
914–925. doi:10.1111/j.1600-0587.2013.00056.x
Newsome, T. M., Stephens, D., Ballard, G. A., Dickman, C. R., and Fleming,
P. J. (2013b). Genetic profile of dingoes (Canis lupus dingo) and free-
roaming domestic dogs (C. l. familiaris) in the Tanami Desert. Australian
Wildlife Research 40, 196–206. doi:10.1071/WR12128
Newsome, T. M., Ballard, G. A., Crowther, M. S., Fleming, P. J., and
Dickman, C. R. (2014). Dietary niche overlap of free-roaming dingoes
and domestic dogs: the role of human-provided food. Journal of
Mammalogy 95, 392–403. doi:10.1644/13-MAMM-A-145.1
Person, D., and Russell, A. (2009). Reproduction and den site section
by wolves in a disturbed landscape. Northwest Science 83, 211–224.
doi:10.3955/046.083.0305
Smith, B. (2015). Biology and behaviour of the dingo. In ‘The Dingo
Debate: Origins, Behaviour and Conservation’. (Ed. B. Smith.)
pp. 25–53. (CSIRO Publishing: Melbourne.)
Thomson, P. (1992a). The behavioural ecology of dingoes in north-western
Australia. II. Activity patterns, breeding season and pup rearing. Wildlife
Research 19, 519–530. doi:10.1071/WR9920519
Thomson, P. (1992b). The behavioural ecology of dingoes in north-western
Australia. III. Hunting and feeding behaviour, and diet. Wildlife
Research 19, 531–541. doi:10.1071/WR9920531
Thomson, P. (1992c). The behavioural ecology of dingoes in north-western
Australia. IV. Social and spatial organisation, and movements. Wildlife
Research 19, 543–563. doi:10.1071/WR9920543
HAustralian Mammalogy B. Smith and A.-L. Vague
www.publish.csiro.au/journals/am