Top-dogs and under-dogs: Competition between dogs and sympatric carnivores

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DOI: 10.1093/acprof:osobl/9780199663217.003.0003
In book: Free-Ranging Dogs and Wildlife Conservation, Chapter: 3, Publisher: Oxford University Press, Editors: Matthew E. Gompper, pp.69-93
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Abstract
This chapter examines the interaction of dogs with other predators. It studies the competitive dynamic existing between dogs and other carnivores. It also assesses the implications of these reactions to the conservation of native carnivore species.
Free-Ranging Dogs and Wildlife Conservation. Edited by Matthew E. Gompper
© Oxford University Press 2014. Published 2014 by Oxford University Press.
dogs with sympatric carnivores will depend on
their position in the interference hierarchy, with
dogs being either at the receiving or giving end
of aggressive interactions, or both.
2. Dependence on human-derived food: Interfer-
ence competition may be seen ultimately as a
function of food acquisition. Although most dog
populations are heavily dependent on human-
derived food (HDF), this is variable depend-
ing on their degree of association with humans.
Dogs in urban areas are almost completely de-
pendent on HDF, whereas at the other extreme,
wild dogs (such as the dingo) and feral dogs are
fully dependent on wild-caught food ( Vanak and
Gompper, 2009b ). Thus, it can be assumed that
competition with wild carnivores is likely high-
est in areas where dependence on wild resources
is highest.
3. Population density: The competitive effects of
dogs can also be in uenced by their density and
their tendency to form packs. Even without hu-
man involvement, packs of dogs are capable of
overpowering not just herbivores several times
their size ( Corbett, 1995 ), but also carnivores
such as foxes, coyotes, and jackals ( Vanak and
Gompper, 2009b ; Van Sittert, 1998 ). Furthermore,
numerical superiority can improve competitive-
ness in obtaining resources such as carcasses.
Thus, competitive ability may be a function of
density at the local scale.
3.1 Introduction
In many natural and semi-natural systems world-
wide, dogs ( Canis familiaris ) are an integral part of
the carnivore community (Gompper, Chapter 1 ).
Until recently, the role of dogs as mid-sized mem-
bers of the carnivore guild, and how they shape
the structure of native carnivore communities, has
garnered little attention. Of late, however, a suite
of studies across the world has highlighted several
key aspects of the competitive dynamics that occur
between dogs and sympatric members of the car-
nivore community ( Table 3.1 ). These studies have
demonstrated that dogs often compete with native
carnivores for food, and that smaller carnivores
may react to dogs as they would to any other mid-
sized predator: with increased vigilance, lowered
food intake, avoidance of dog-dominated habitat
types, or complete spatial separation. These studies
have also shown that the types and intensity of in-
teractions between dogs and subordinate predators
are likely in uenced by several factors, including,
but not limited to:
1. Relative position within the native carnivore
community: In several areas dogs are the largest
mammalian carnivore and hence assume the role
of the apex predator (e.g., Australasia), whereas
in others they may be part of a guild of carnivores
where they occupy a mesopredator position (e.g.,
India, Africa). The nature of the interactions of
CHAPTER 3
Top-dogs and under-dogs:
competition between dogs
and sympatric carnivores
Abi Tamim Vanak , Christopher R. Dickman , Eduardo A. Silva-Rodriguez ,
James R.A. Butler , and Euan G. Ritchie
04-Gompper-Chap03.indd 6904-Gompper-Chap03.indd 69 14/09/13 9:34 AM14/09/13 9:34 AM
70 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
Table 3.1 Summary of studies that examine or report the competitive effects of dogs on sympatric carnivores.
Country Predator guild*, besides
dog
Interactions
studied
Effect of dogs Reference
Australia Large marsupials Dog (dingo)
thylacine and
Tasmanian devil
Direct killing and possibly more effi -
cient resource exploitation, resulting
in extinction of both marsupials on
the Australian mainland
§
Corbett 1995 ; Glen and
Dickman 2005 ; Wroe et al.
2007 ; Fillios et al. 2012
Australia Invasive red fox, feral cat,
and small to mid-size
native predators
Dog (dingo) fox
and cat
Killing, harassment, spatial
interference (fox, cat) with
corresponding benefi t to native
species via mesopredator release
#
Corbett 1995 ; Glen and Dick-
man 2005 ; Mitchell and Banks
2005 ; Johnson and VanDerWal
2009 ; Letnic et al. 2009 ; 2011 ;
2012 ; Purcell 2010 ; Wallach
et al. 2010 ; Brawata and
Neeman 2011 ; Kennedy et al.
2012 ; Moseby et al. 2012
Brazil Maned wolf, crab-eating fox,
puma, South American coati,
crab-eating raccoon, tayra
Dog Maned wolf Interference on maned wolf, no
evidence of effects on crab-eating
raccoon §
Lacerda et al. 2009
Brazil Crab-eating fox, South
American coati, crab-eating
raccoon, tayra, lesser grison,
oncilla, margay
Dog Crab-eating
fox and coati
Evidence of negative association
with crab-eating fox and coati, no
evidence of effects on tayra and
small cats
§
Espartosa 2009
Chile Puma, chilla fox, guigna,
Molina’s hog-nosed skunk,
American mink (invasive)
Dog Chilla fox Killing, harassment, spatial
interference
#
Silva-Rodriguez et al. 2010a
Ethiopia Ethiopian wolf Dog → ← Ethiopian
wolf
Outcome of interactions depended
on numerical superiority
#
Atickem et al. 2010
India Wolf, golden jackal, jungle
cat, Indian fox
Dog Indian fox Killing, harassment, spatial
interference
#
Vanak et al. 2009 ; Vanak and
Gompper 2010
India Striped hyena, wolf, jungle
cat, Indian fox
Dog golden
jackal
Dominance at carcasses § Aiyadurai and Jhala 2006
Madagascar Fossa, small-toothed civet,
Malagasy civet, ring-tailed
mongoose, broad-striped
mongoose, small Indian
civet, domestic cat
Dog Fossa and
ring-tailed mongoose
Fossa nearly absent from sites
with high dog occupancy. Capture
rates of fossa decreased as capture
rates of dogs increased. Ring-tailed
mongoose changed activity patterns
when dogs and the invasive small
Indian civet were present. No effects
reported for other three species
analyzed §
Gerber et al. 2012 ; Barcala
2009
Poland Red fox, domestic cat Dog Red fox, cat Negative association between
occurrence of dogs with cats and
red foxes
§
Krauze-Gryz et al. 2012
Spain Red fox, badger, domestic
cat, Egyptian mongoose,
Dog Badger Spatial exclusion § Revilla et al. 2001
United Kingdom Red fox Dog Red fox Killing of adults and pups Harris 1981
Zimbabwe Lion, spotted hyena,
leopard, black-backed
jackal, side-striped jackal
Lion, spotted hyena,
leopard Dog
jackals
Dominance over jackals at carcasses
on wildlife reserve peripheries;
spatial exclusion of jackals (in
communal lands and on commercial
farmland peripheries) #
Butler 1998 ; Butler and
Bingham 2000 ; Butler and du
Toit 2002 ; Butler et al. 2004
* Arranged in order of decreasing body size,
# Direct evidence,
§ Indirect or correlative evidence. Arrows indicate direction of dominant interactions.
04-Gompper-Chap03.indd 7004-Gompper-Chap03.indd 70 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 71
3.2 Dogs as interference competitors
Predators that share common resources will often
show potential to compete with each other, but
even if resources are abundant and overlap is mini-
mal, interference competition may still occur via
direct aggression and intraguild killing or via cues
that drive spatial segregation ( Box 3.1 ). Interfer-
ence competition between mammalian carnivores
is well documented, and it is expected that dogs
as abundant mid-sized carnivores will be either re-
cipients of top-down interference or will be the ag-
gressors. However, the competitive dynamics that
occur between dogs and sympatric carnivores can
be expected to vary considerably across the world
as a function of related human densities, land use
types, and the place of dogs in the native carnivore
community.
3.2.1 Where dogs are top-predators
In many regions of the world, dogs have assumed
the role of top-predators for a variety of reasons: the
absence of larger mammalian carnivores, the exter-
mination of large carnivores either by humans or
other factors, or the restriction of larger carnivores
to non-human dominated landscapes ( Gompper
and Vanak, 2008 ). Some potential examples of the
competitive ability of dogs as top-predators come
4. Ranging behavior: The ability of dogs to range
widely into natural habitats increases the poten-
tial for them to compete with sympatric carnivores
not just at the periphery of human settlements
but also in native habitats. Vanak and Gompper
( 2009b ) proposed a conceptual model wherein
the competitive effects of dogs were affected not
just by their densities, but also by their ranging
behavior ( Figure 3.1 ). According to this model,
the maximum impact on sympatric carnivores is
expected from a high-density dog population that
also exhibits wide-ranging behavior.
Other than in Australasia, few dog populations are
truly independent of humans ( Vanak and Gompper
2009b ). Ultimately, their close relationship with hu-
man society strongly in uences all the above fac-
tors. Whether dogs are herding dogs, guard dogs,
village dogs, farm dogs, or simply household pets
determines in large part their ranging behavior,
population density, and feeding habits. Close as-
sociation with human habitation may also provide
refugia from top-down interference competition or
predation in systems where larger predators exist
(Butler et al., Chapter 5 ).
Here, we review the literature to examine compet-
itive dynamics between dogs and sympatric carni-
vores as a function of their position in the carnivore
community and the type of competition (see Box 3.1 ).
Population density
Ranging behavior
High Low
Low
High
Low High
Risk to sympatric carnivores
Figure 3.1 Conceptual model of the
effect of dog population density and
ranging behavior on the risk dogs may
pose as interference competitors to
sympatric native carnivores. Adapted from
Vanak and Gompper ( 2009b ) . Reproduced
with permission from John Wiley & Sons.
04-Gompper-Chap03.indd 7104-Gompper-Chap03.indd 71 14/09/13 9:34 AM14/09/13 9:34 AM
72 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
Box 3.1 Competition in its various forms
Within the carnivore guild, sympatric species compete fi ercely
with each other and this interaction manifests as either a form
of: (1) exploitative competition, whereby differences in acqui-
sition of limited resources determine the competitive out-
come ( Figure 3.2a ); or (2) interference competition, whereby
direct interactions such as spatial exclusion, harassment, or
even intraguild killing determine the outcome ( Figure 3.2b ).
Exploitative competition in carnivores has not been directly
demonstrated but can be inferred, especially for species that
kleptoparasitize the kills of others. Interference competition
and intraguild killing, however, are common among carni-
vores and there are many well-documented examples of
these interactions.
Interference competition
Mechanisms of interference competition usually follow an al-
lometric relationship, with larger carnivores directly dominating
smaller ones. Smaller species thus have a strong imperative to
minimize interference interactions, especially when there are
potentially lethal consequences ( Creel et al., 2001 ; Palomares
and Caro, 1999 ; Ritchie and Johnson, 2009 ). Studies that ex-
amine single or multiple features of interference competition
among carnivores are numerous and have been reviewed ex-
tensively in the literature ( Creel et al., 2001 ; Glen and Dickman,
2005 ; Ritchie and Johnson, 2009 ; Ritchie et al., 2012 ; Roemer
et al., 2009 ). Here we highlight some of the patterns.
In general, interference competition can affect subor-
dinate competitors in several ways: by limiting spatial dis-
tributions ( Creel et al., 2001 ; Ritchie and Johnson, 2009 ),
constraining habitat selection ( Mitchell and Banks, 2005 ;
Vanak and Gompper, 2010 ; Fisher et al., 2013), or reducing
prey encounter rates ( Atwood and Gese, 2008 ; Creel et al.,
2001 ; Palomares et al., 1996 ; Vanak et al., 2009 ). In response,
subordinate competitors use a suite of spatial and temporal
behavioral strategies to minimize encounters with the domi-
nant competitor(s). These include avoiding range overlap
altogether, or modifying range use within overlapping home
ranges (e.g., Berger and Gese, 2007 ; Crabtree and Sheldon,
1999 ; Elmhagen et al., 2002 ; Fedriani et al., 2000 ; Gosselink
et al., 2003 ; Tannerfeldt et al., 2002 ). In these cases, inter-
ference competition results in non-overlapping territories, in
favor of the dominant competitor.
There are also more subtle spatial and temporal behav-
ioral choices that subordinate competitors make to avoid
interference competition. African hunting dogs (
Lycaon
pictus
) avoid prey-rich habitat due to the presence of li-
ons (
Panthera leo
) ( Creel et al., 2001 ), thereby incurring
a reduction in hunting effi ciency. After the reintroduction
of wolves (
C. lupus
) into Yellowstone National Park, USA,
coyotes (
C. latrans
) reduced foraging and increased vigilance
when they were in wolf territories ( Switalski, 2003 ). Kit foxes
(
Vulpes macrotis
) are known to use several anti-predatory
tactics, such as multiple diurnal den use and habitat parti-
tioning, to avoid interference competition with coyotes ( Nel-
son et al., 2007 ). These examples demonstrate that the mere
presence of a dominant competitor results in risk aversion
that is manifested by the reduced use of areas that would
otherwise be selected for. Ultimately, interference competi-
tion can result in reduced individual fi tness and population
sizes ( Ritchie and Johnson, 2009 ).
+
+
(a)
–/0
(b)
+ +
(c)
Figure 3.2 Illustrations of exploitative
(a), interference (b), and apparent
competition (c). Solid arrows denote direct
effects, dashed arrows indicate indirect
effects, and signs indicate effect on
species. Adapted from Glen and Dickman
( 2005 ). Reproduced with permission from
John Wiley & Sons.
continued
04-Gompper-Chap03.indd 7204-Gompper-Chap03.indd 72 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 73
Wroe et al., 2007 ). The evidence is ostensibly com-
pelling. In Tasmania, thylacines (15–35 kg) report-
edly hunted kangaroos and wallabies in open forest
and woodland areas, pursuing their quarry singly
or in pairs at night ( Paddle, 2002 ). They may also
have hunted a range of smaller prey ( Attard et al.,
2011 ; Wroe et al., 2007 ). Tasmanian devils (5–15 kg)
occupy similar habitats and also pursue wallabies
and smaller mammals, but include much scavenged
material in their diet ( Jones and Barmuta, 1998 ). Both
species probably would have overlapped markedly
in these aspects of their resource use with the dingo.
from Australia. Prior to the arrival of the dingo in
Australia around 4,000 years ago, the largest mam-
malian predators were the thylacine ( Thylacinus
cynocephalus ) and the Tasmanian devil ( Sarcophilus
harrisii ). Both of these marsupial predators disap-
peared from the Australian mainland after dingoes
had become established, the thylacine around 3,500
years ago and the devil within the last millennium
( Corbett, 1995 ). As both species persisted in the ab-
sence of dingoes in Tasmania, it has been commonly
assumed that their demise on the mainland was
caused by competition from the dingo ( Corbett, 1995 ;
Box 3.1
Continued
Exploitative competition
Exploitative competition occurs when species share the
same limited resources, and one species can potentially out-
compete the other either through numerical or behavioral
superiority in acquiring this shared resource ( Petren and
Case, 1996 ). Demonstrating the occurrence of exploitative
competition between two species requires a number of fac-
tors to fi rst be established: (1) reduced survivorship or re-
production as a function of the limited and shared resource;
(2) reduced access to the shared resource; and (3) a lack of
direct interference ( Petren and Case, 1996 ). Because of this,
exploitative competition among carnivores has rarely been
demonstrated even though there is extensive evidence of
dietary overlap between species. Indeed, exploitative com-
petition has only been experimentally demonstrated in a
few plants and animals ( Bonaccorso et al., 2007 ; Dorchin,
2006 ; Smallegange et al., 2006 ). Nonetheless, the existence
of exploitative competition is a common assumption among
carnivore ecologists ( Jhala and Giles, 1991 ; Johnson et al.,
1996 ; Sillero-Zubiri and Gottelli, 1995 ).
Apparent competition
A third kind of interspecifi c interaction may also function to
explain patterns of carnivore spatial distribution: apparent
competition. While interference and exploitation competi-
tion are perceived as a function of resource limitation, the
outcome of apparent competition results from a third factor
indirectly mediating the distribution of apparently competing
species ( Figure 3.2c ) ( Holt, 1977 ; Holt and Lawton, 1994 ;
Price et al., 1998 ). For example, populations of Sierra Nevada
bighorn sheep (
Ovis canadensis californiana
) declined due
to predation from mountain lion (
Puma concolor
), whose
numbers were primarily sustained by the more abundant
mule deer (
Odocoileus hemionus
). Hence, without factoring
in predation by mountain lion, the decline in bighorn sheep
appeared to be due to competition with mule deer ( Johnson
et al., 2013 ). Similarly, the critically endangered island fox
(
Urocyon littoralis
) faced near extinction due to predation by
golden eagles (
Aquila chrysaetos
), whose unnaturally large
populations were in turn sustained by invasive feral pigs (
Sus
scrofa
) ( Roemer et al., 2001 , 2002 ). Apparent competition
among carnivores may result from one carnivore subsidizing
the presence of another carnivore, and thus also putatively
reducing the density of other sympatric carnivore species.
However, apparent competition is more likely to result from
shared parasites than from shared predators. Shared para-
sites may be an important driver in structuring assemblages
of species ( Rushton et al., 2000 ; Tompkins et al., 2000 ,
2003 ). This kind of interspecifi c interaction is a form of ap-
parent competition because the parasite–host interactions
result in the appearance of a competitive dynamic ( Holt,
1977 ; Holt and Pickering, 1985 ; Price et al., 1998 ; Tompkins
et al., 2000 ). Apparent competition has been documented
in many taxa, ranging from bacteria and insects to birds
and large mammalian predator–prey communities. In gen-
eral, the rarer or more susceptible host species is reduced or
eliminated as the parasite increases in the less susceptible
or more abundant reservoir host (e.g., Morris et al., 2004 ;
Power and Mitchell, 2004 ; Tompkins et al., 2000 ). Appar-
ent competition via shared parasites may ultimately turn out
to be a common and important component of community
structure ( Hatcher et al., 2006 ; Holt and Dobson, 2006 ) but
has only rarely been explored as such in carnivore communi-
ties ( Roemer et al., 2009 ).
04-Gompper-Chap03.indd 7304-Gompper-Chap03.indd 73 14/09/13 9:34 AM14/09/13 9:34 AM
74 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
( Fillios et al., 2012 ). As this suite of changes affected
the Australian mainland (but Tasmania to a much
lesser extent), the arrival of the dingo would have
had an additive impact on the mainland popula-
tions of the thylacine and devil and may perhaps
have delivered the competitive coup de grace . The
longer tenure of devils in the presence of dingoes on
the Australian mainland may have arisen from their
ability to exploit carrion and small prey that formed
a more minor part of the diet of the dingo; that is,
there was perhaps more dietary niche separation and
hence less competition between devils and dingoes
than between dingoes and thylacines ( Corbett, 1995 ).
As the apex mammalian predator in Australia,
dingoes could be expected to dominate in direct
encounters with most other predators; moderate to
large crocodiles would have ascendancy over them,
although their eggs and young still may be vulner-
able ( Somaweera et al., 2011 ). No studies have yet
addressed whether sympatric predators recognize
and avoid cues of the presence of the dingo. There
is some evidence that native prey species can distin-
guish the odor or other cues left by dingoes and do-
mestic dogs and reduce their activity after exposure
( Carthey and Banks, 2012 ; Parsons and Blumstein,
2010 ); native predators plausibly also would have
experienced selection to reduce their risk of an en-
counter with dingoes.
In many parts of the world with high human pop-
ulation densities, native predators have either been
Although dingoes are smaller than the Tasmanian
thylacine, at 12–22 kg, their ability to hunt in packs
may have given them a competitive edge over the
marsupial predators and suppressed their popula-
tions via either extreme interference competition or
superior exploitation of shared prey ( Letnic et al.,
2012 ). Recent evidence further indicates that main-
land thylacines were smaller than their counterparts
in Tasmania and that mainland females were con-
siderably smaller than dingoes ( Fillios et al., 2012 ).
This potentially would have placed mainland thy-
lacines under great pressure, especially from direct
interference interactions.
Despite the elegance of this interpretation, the
arrival of the dingo coincided with several envi-
ronmental changes that may have been more det-
rimental to the marsupials, making it dif cult to
resolve the competitive impact of the dingo on its
own. The mid Holocene experienced intense cli-
matic changes as the continent recovered from the
extremes of the last glacial maximum ( Brown, 2006 ).
There was also an increase in the human popula-
tion and in peoples’ hunting ef ciency due to the
adoption of technologies, such as edge-ground
and hafted stone tools, that improved consider-
ably on previous weaponry ( Johnson and Wroe,
2003 ; Letnic et al., 2012 ). Coincident with (and per-
haps caused by) these changes, archeological re-
cords suggest that people exploited smaller-bodied
prey progressively from the mid to late Holocene
Box 3.2 Old dogs and new dogs: competition between dingoes and recent dogs
As a general rule, the form of competition that occurs be-
tween two species will be the same form of competition that
is apparent between conspecifi cs. Thus, we might expect
that interference will be the dominant form of competition
between different groups of dogs, with exploitation play-
ing no, or a very minor, role. Studies of wild animals gen-
erally bear out this expectation: pairs or packs of dingoes,
for example, use howling and scent-marking to defend the
boundaries of their territories, and chase intruders if they are
detected ( Purcell, 2010 ). There is no evidence for exploita-
tive interactions between packs.
But what if dogs differ from each other in subtle ways?
Dingoes and recently introduced domestic dogs co-occur
throughout much of Australia and differ in aspects of their
size, skull morphology, coloration, behavior, and seasonality
of reproduction ( Corbett, 1995 ). Dingoes occur in most parts
of the continent except where they are heavily persecuted in
the south-east and south-west, whereas wild domestic dogs
tend to be most active near townships and settlements. There
is some evidence that wild domestic dogs seldom successfully
breach the territorial boundaries established by dingo packs,
and fare poorly due to interference ( Corbett, 1995 ). Conversely,
continued
04-Gompper-Chap03.indd 7404-Gompper-Chap03.indd 74 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 75
exposed to dogs, foxes reduced food consumption
by as much as 70% ( Figure 3.3a ) and increased time
spent in vigilance behavior 15-fold ( Figure 3.3b ;
Vanak et al., 2009 ). Thus, at the  ne scale, Indian fox-
es showed a strong foraging-vigilance trade-off, but
risk-averse behaviors were also seen at larger spatial
scales. The odds of foxes using an area decreased
rapidly the closer that area was to a dog-dominated
habitat. This happened even if these areas were as-
sociated with a higher abundance of their main food
source ( Vanak and Gompper, 2010 ).
Similar spatial avoidance in response to interfer-
ence competition is also observed in other parts of
the world. In the United Kingdom, the distribution
of red foxes ( V. vulpes ) was negatively affected by the
presence of high densities of street dogs. These dogs
were not only the second most important cause of
mortality of cubs, with as much as 15% of cub mor-
tality attributed to dogs, but also killed adult foxes
on occasion ( Harris, 1981 ). In South America, three
different studies suggest that dogs have negative ef-
fects on the occurrence of canids such as the maned
wolf ( Chrysocyon brachyurus ), chilla fox ( Lycalopex
exterminated or occur at very low densities. Here,
free-ranging dogs have become the de facto top-
predator, and this is re ected in their interactions
with the remaining native carnivores. One of the most
comprehensive examples of competitive dynamics
between dogs and sympatric carnivores comes from
the dry grasslands of Maharashtra, India, where
dogs interact with native Indian foxes ( V. bengalen-
sis ). Here, dogs are not the largest predator, as the
Indian wolf ( C. l. pallipes ) occupies that spot. Howev-
er, wolves occur at very low densities ( Habib, 2007 ),
while dogs occur at very high densities (24 dogs/
sq. km; Vanak, 2008 ; Vanak and Gompper, 2010 ).
Thus, as the most abundant mid-sized carnivore in
the landscape, dogs are dominant. Dogs and Indian
foxes have low dietary overlap and thus do not ap-
pear to compete directly for food. This is due mainly
to the fact that dogs subsist almost entirely on HDF
such as garbage, human feces, direct feeding, and
carrion, whereas foxes subsist almost entirely on ro-
dents, fruit, and invertebrates ( Vanak and Gompper,
2009 a). Yet, Indian foxes react to dogs as they would
to a dominant carnivore. When experimentally
Box 3.2
Continued
large breeds of domestic dog, such as the Maremma, are be-
ing used increasingly on rural properties to protect fl ocks of
sheep from the ravages of dingoes. Termed livestock guard-
ian dogs (LGDs), these large and aggressive animals are al-
lowed to bond with fl ocks from an early age and are provided
with all their food and shelter needs by their human owners.
Once trained, they can be very effective in using aggressive
interference behaviors to repel dingoes. In one survey of 150
livestock producers, two thirds of respondents indicated that
predation from dingoes and other wild dogs ceased after
LGDs were put in service; another 30% reported that preda-
tion on livestock decreased ( van Bommel and Johnson, 2012 ).
In these situations, LGDs are essentially the guardians of their
own human-defi ned territories (usually fences and paddock
boundaries) and effectively repel incursions by dingoes via
direct interference. Without the continuous resource-subsidy
from humans it is not clear whether LGDs would always pre-
vail over dingoes, although interference would almost cer-
tainly be the mechanism of competition in any encounters.
Over the long term, any competitive superiority of dingoes
over wild domestic dogs may not be suffi cient for dingoes
to survive in ‘pure’ form. Another kind of interaction—
hybridization—may doom them. Although wild domestic
dogs seldom gain access to dingo packs, they can do so
more easily after packs have been fractured by misguided
human attempts to control dingo numbers. Such attempts,
using poisons, trapping, or shooting, often remove the
dominant animals from a pack and may loosen the social
and reproductive controls on remaining animals suffi ciently
that wild dogs then can gain access to the packs ( Allen and
Gonzalez, 1998 ; Purcell, 2010 ). If breeding is successful,
hybridization ensues. Recent studies suggest that dingoes
in remote regions of Australia have suffered little hybridiza-
tion, with up to 90% of animals retaining the ‘pure’ dingo
genotype (Newsome et al., 2013), whereas only 15% or less
may be ‘pure’ in more heavily settled and rural areas where
dingoes are controlled and wild domestic dogs are more
prevalent ( Elledge et al., 2009 ). Hybridization is not usually
seen as a form of competition. However, the inexorable sup-
planting of dingo by dog genes in the present example gives
the process the appearance of a strong and highly asym-
metrical form of competition.
04-Gompper-Chap03.indd 7504-Gompper-Chap03.indd 75 14/09/13 9:34 AM14/09/13 9:34 AM
76 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
(see Box 3.3 ). For example, in Spain the presence
of dogs was inversely correlated with an indicator
of Eurasian badger ( Meles meles ) presence ( Revilla
et al., 2001 ). In Madagascar, the fossa ( Cryptoprocta
ferox ) is nearly absent from areas with high dog oc-
cupancy ( Gerber et al., 2012 ) and trapping rates of
this species decrease as dog trapping rates increase
( Barcala, 2009 ; see Box 3.4 ). Similarly, the highest
densities of the endangered narrow-striped mon-
goose ( Mungotictis decemlineata ) occur in areas
where neither dogs nor people are detected and,
in one area, sightings of this species increased after
dog removal ( Woolaver et al., 2006 ). These comple-
mentary distribution patterns are likely to be com-
mon throughout the world, especially in countries
where dog densities are largely unregulated in
rural areas ( Boitani et al., 1995 ; Krauze-Gryz and
Gryz, 2009 ; Slater et al., 2008 ; Vanak and Gompper,
2010 ).
griseus ), and crab-eating fox ( Cerdocyon thous ) ( Es-
partosa, 2009 ; Lacerda et al., 2009 ; Silva-Rodríguez
et al. 2010a ). Two of these studies also addressed
the effects of dogs on non-canid carnivores such
as South American coati ( Nasua nasua ), crab-eating
raccoon ( Procyon cancrivorus ), tayra ( Eira barbara ),
and small cats ( Leopardus spp.). Among these spe-
cies, and with the sole exception of the coati ( Es-
partosa, 2009 ), there was no evidence of potentially
negative effects of dogs ( Espartosa, 2009 ; Lacerda et
al., 2009 ). The fact that canids seem to show stronger
responses to dogs than other carnivores is not sur-
prising as interference competition and intraguild
killing should be more intense between members of
the same family ( Donadio and Buskirk, 2006 ).
Because of the scarcity of directed studies on
competition between dogs and sympatric carni-
vores, the competitive effect of dogs can sometimes
only be inferred without a direct link to causality
(a)
(b)
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
Neutral Live dog Odors
Proportion food eaten at trays
*
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
Non-vigilance Vigilance At food trays
Proportion of time spent
a
b
a
a
b
a
a
b,c
a,c
Neutral
Live dog
Odors
Figure 3.3 Indian foxes, when
exposed to the presence of a live dog
at experimentally placed food trays, (a)
reduced the proportion of food eaten by
~70% compared to a control tray, and (b)
increased time spent in vigilance behaviors
by 15-fold when compared to trials when
only dog odor or a control was placed.
The asterisk and letters denote signifi cant
difference (Fisher’s protected least squares
difference
P
<0.05) among trials for
each behavior and error bars represent
mean ± SE,
n
= 7 dens. Reprinted from
Vanak et al. ( 2009 ) with permission from
Springer.
04-Gompper-Chap03.indd 7604-Gompper-Chap03.indd 76 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 77
Box 3.3 Competition between dingoes and introduced predators
In contrast to the relative paucity of studies invoking com-
petition between dingoes and native predators, a substan-
tial body of work has focused on interactions between
dingoes and the more-recently introduced red fox and feral
cat (
Felis catus
). Both predators show varying degrees of
dietary overlap with the dingo, although their small size
(fox 4–8 kg; cat 3–6 kg) means that they generally hunt
smaller prey than their larger counterpart ( Cupples et al.,
2011 ; Glen et al., 2011 ). All three species are broadly sym-
patric over much of Australia and often co-occur in the
same habitat, thus increasing the opportunity for competi-
tion. Evidence that competition does indeed occur is de-
rived largely from inverse spatial patterns in the species’
abundances. In the case of the fox, several studies have
shown that fox numbers and/or activity are suppressed
in the presence of the dingo at local, regional, and conti-
nental scales ( Johnson and VanDerWal, 2009 ; Letnic et al.,
2009 , 2012 ; Mitchell and Banks, 2005 ; Wallach et al.,
2010 ). In general these studies also show that where din-
goes are abundant, foxes are rare, but where dingoes are
scarce or absent, foxes can achieve much higher numbers
( Figure 3.4 ). In the case of the feral cat, evidence for com-
petition is more tenuous. Several studies show negative
associations between the numbers of dingoes and cats
( Brawata and Neeman, 2011 ; Kennedy et al., 2012 ; Pur-
cell, 2010 ; Wallach et al., 2010 ), but others show weak
patterns or no relationship ( Letnic et al., 2009 ; Newsome,
2001 ). In contrast to the red fox, which is largely active on
the ground, feral cats can climb trees and thus may be bet-
ter able to avoid direct contact with dingoes. For example,
in very barren areas of the Channel Country in western
Queensland, the primary refuge sites are tree-lined ripar-
ian strips that allow cats to escape ground-active dingoes
( Pettigrew, 1993 ).
450
400
350
300
250
200
150
100
50
0
(a)
0 200 400 600 800
Dingo
bounties
(b)
0 50 100 150 200
Dingo
bounties
4000
3500
3000
2500
2000
1500
1000
500
0
Fox
bounties
Fox
bounties
Figure 3.4 Numerical relationship
between the dingo and red fox,
based on bounty records collected (a)
in Queensland in the fi nancial year
1951–52 and (b) in Western Australia in
the years 1947–52. Each dot represents
the number of bounties paid within
100 × 100 km grid cells in each state
over the bounty collection period. Solid
lines show regressions for the 90%
quantile of fox abundance against dingo
abundance, light dotted lines show
signifi cant (
P
<0.05) least squares linear
regressions, and the heavy dotted line
in (a) indicates a signifi cant (
P
<0.05)
least squares logarithmic regression.
Reprinted from Letnic et al. ( 2011 ) with
permission from John Wiley & Sons.
continued
04-Gompper-Chap03.indd 7704-Gompper-Chap03.indd 77 14/09/13 9:34 AM14/09/13 9:34 AM
78 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
suppressed by the presence of larger competitors
( Crooks and Soulé, 1999 ). Thus, it would be expect-
ed that in areas where larger carnivores are present,
dogs would be negatively affected. However, be-
cause of the lack of directed studies in such systems,
3.2.2 Dogs as mesocarnivores
In several ecosystems across the world, dogs are
sympatric with larger carnivores such as the wolf,
leopard ( Panthera pardus ), tiger ( P. tigris ), and
bears. In natural systems, mesocarnivores are often
Box 3.3
Continued
The form of interaction between dingoes and the two
smaller predators is almost certainly extreme interference
competition. This is suggested by the fact that red fox and
feral cat remains are found frequently but only at very low
levels in scat or stomach samples of the dingo ( Allen and
Leung, 2012 ; Letnic et al., 2012 ), and observations that both
the smaller predators will fl ee if an encounter with a dingo
is imminent ( Corbett, 1995 ; Purcell, 2010 ). However, the
most compelling evidence for extreme interference comes
from observations of dingoes killing but not eating both
foxes and feral cats in a large (37 km
2 ) enclosure in treeless
habitat in South Australia ( Moseby et al. 2012 ). In this latter
study, data derived from GPS collars confi rmed that dingoes
returned to the carcasses of foxes and cats that they had
killed for several hours after the killing, with some dingoes
returning again and again over the ensuing days. It is pos-
sible that these frequent return visits were to ensure that
the smaller predators no longer posed a competitive threat.
Box 3.4 Landscapes of fear: spatial and temporal considerations
There is increasing recognition that predators, including
dogs, may infl uence their prey in subtle ways that are not
readily apparent by simply comparing measures of abun-
dance of pairs of interacting species ( Creel and Christianson,
2008 ; Ritchie and Johnson, 2009 ). This is important, be-
cause signifi cant effects of dogs on other species may have
been overlooked or dismissed previously due to a focus on
abundance patterns only ( Johnson and VanDerWal, 2009 ).
Recent studies from two vastly separated regions (Australia
and Madagascar) and with different predator communities
serve to illustrate this point.
In Australia, dingoes are often killed to minimize their at-
tacks on livestock, and the impact this has on their abun-
dance and behavior, as well as that in turn on a sympatric
mesopredator, the feral cat, was examined using remote
camera surveys across nine pairs of large Australian range-
land (cattle) properties ( Brook et al., 2012 ). Dingo abun-
dance was generally reduced on properties where dingoes
were controlled, but most interestingly dingo behavior var-
ied between sites where dingoes were controlled and where
they were not. Uncontrolled dingo populations were crepus-
cular, similar to their major prey (kangaroos). In populations
subject to control, dingoes were less active around dusk,
with activity concentrated in the period immediately before
dawn. Shifts in feral cat abundance were inversely related
to corresponding shifts in dingo abundance, but importantly
there was also a negative relationship between predator
visitation rates at individual camera stations, indicating that
cats avoided areas where dingoes were most active, and
that reduced activity by dingoes at dusk was associated with
higher cat activity at dusk. This suggests that dingo control
may not only lead to more feral cats (mesopredator release),
but also allows cats to optimize their hunting when dingoes
are less active ( Brook et al., 2012 ).
In Madagascar, as part of an overall study examining
activity patterns of carnivores, the authors found that the
fossa selectively used crepuscular hours, but was cathem-
eral overall ( Gerber et al., 2012 ). Fossas were conspicuously
absent from sites where dogs were most abundant and ac-
tive throughout the diel cycle, and the ring-tailed mongoose
(
Galidia elegans
), another native, but smaller-bodied preda-
tor, shifted its activity away from its preferred periods in the
presence of dogs. Together, these studies highlight that indi-
cators other than abundance, such as spatial and temporal
behavioral patterns, should also be considered when deter-
mining a predator’s ability to affect trophic levels. However,
abundance still remains an important consideration, as dogs
have even been known to exclude larger-bodied predators,
such as wolves, from accessing food; despite their smaller
body-size as compared to wolves, dogs overcame this by be-
ing more aggressive and occurring in higher numbers than
wolves ( Boitani et al., 1995 ).
04-Gompper-Chap03.indd 7804-Gompper-Chap03.indd 78 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 79
production, dogs occur at very low densities due
to the lack of human settlement. As a consequence
side-striped and black-backed jackals ( C. mesomelas )
have become the most numerous native carnivore
through ‘mesopredator release,’ occurring at high
densities relative to natural conditions ( Cumming,
1982 ; Bingham et al. 1999 ). Hence within commu-
nal lands and where they border commercial farm-
land, relationships within the carnivore guild are
limited to competitive interactions between dogs
and jackals.
Both jackal species have omnivorous and oppor-
tunistic feeding habits in Africa ( Skinner and Chim-
imba, 2005 ), and will ef ciently scavenge carrion
( Richardson, 1980 ) and human waste ( Bothma et al.,
1984 ), potentially overlapping the dietary niche of
dogs. Consequently the ecological relationship be-
tween dogs and jackals is characterized by con ict
and intolerance, with dogs dominating jackals in
aggressive encounters due to their larger size (15 kg
versus 8–10 kg for side-striped and 7–8 kg for black-
backed jackals). Carcass experiments showed that
side-striped jackals are attracted to carrion but do
not feed, perhaps avoiding the scavenging dogs
present ( Butler and du Toit, 2002 ; see Section 3.3.1 ).
At higher densities, dogs aggressively exclude jack-
als and jackals may become the subordinate species,
avoiding contact with dogs and hence occurring at
lower densities in communal lands. Commercial
farmland therefore provides a refuge for jackals,
which then ‘spill over’ into the peripheries of com-
munal lands.
Competition between dogs and other carnivores
may not always follow an allometric hierarchical
relationship. In some cases, dogs can out-compete
larger carnivores because of numerical superiority
via frequency-dependent competition. The outcome
of encounters between the endangered Ethiopian
wolf ( C. simensis ) and dogs in the highlands of Ethi-
opia seems to depend on numbers. If wolves out-
number dogs, then the dogs are chased away, and
vice-versa ( Atickem et al., 2010 ). However, the au-
thors concluded that interference competition with
dogs was not a major concern for the conservation
of Ethiopian wolves, but suggested that this could
change in the future as the numbers of people and
dogs grow ( Atickem et al., 2010 ; Laurenson et al.,
1998 ; Sillero-Zubiri and Macdonald, 1997 ).
we can only speculate on whether dogs are subject
to top-down competitive effects from larger carni-
vores. What we do know is that in many systems
dogs form an important part of the diet of larger
carnivores (e.g., leopard; Butler et al., Chapter 5 ).
Thus, it is likely that in such systems dogs will also
display avoidance tactics as shown by other meso-
carnivores. However, dogs may also be insulated
from top-down effects if they can  nd refuge in hu-
man settlements or bene t from human association
while ranging into native habitats.
Studies from southern Africa, where dogs have
the potential to interact with possibly the greatest
number, as well as diversity, of carnivores, well il-
lustrate the complexity of the situations that are
possible ( Butler and du Toit, 2002 ; Butler et al.,
2004 ). Detailed studies of interactions in Zimba-
bwe between dogs and large wild carnivores in
Gokwe Communal Land (GCL) bordering the
Sengwa Wildlife Research Area (SWRA) suggested
that dogs were inef cient and infrequent preda-
tors. Their small group size (mean 1.7) and body
mass (mean 15 kg) was re ected in their diet, which
consisted of scavenged human refuse, feces and
carrion, fruit, and some insects. The vast majority
(87%) of this food was gained independently, with
55% of it derived from carrion, particularly mam-
malian carcasses. Leopards, lions, spotted hyenas
( Crocuta crocuta ), side-striped jackals ( Canis adus-
tus ), and several smaller carnivore species travelled
up to 3 km into GCL ( Butler, 2000 ) while dogs were
observed up to 3 km within the SWRA indepen-
dently of people, and up to 6 km with people, and
consequently were sympatric with these species
in the SWRA–GCL interface. Dogs dominated the
carnivore scavenging guild (see Section 3.3.1 ), but
were preyed upon by lions, leopards, and spotted
hyenas. Thus, inside wildlife reserves, and on their
boundaries with communal lands, dogs were meso-
level carnivores and had diverse relationships with-
in a guild of larger and smaller native mammalian
and avian species.
In most communal land areas and commercial
farmlands where large carnivores are essentially
absent, either due to high human population den-
sities or persecution, dogs are the local apex preda-
tor. However, in commercial farmland typi ed
by intensive cropping and more extensive cattle
04-Gompper-Chap03.indd 7904-Gompper-Chap03.indd 79 14/09/13 9:34 AM14/09/13 9:34 AM
80 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
1948 ). Dogs are also frequent causes of mortality for
the Geoffroy’s cat ( Leopardus geoffroyi ) and culpeo
foxes ( Lycalopex culpaeus ) on certain Argentinean
ranches ( Novaro et al., 2005 ; Pereira et al., 2010 ).
On the other hand, evidence of consumption comes
from analyses of dog diet. One study showed con-
sumption of carnivores such as lesser grison ( Galic-
tis cuja ) and South American coati ( Campos et al.,
2007 ), but it was unclear whether dogs killed these
carnivores or simply scavenged them. Despite these
studies, the quality of the evidence for the impor-
tance of dogs as a source of mortality is mixed. In
fact, most of the evidence available is constituted by
anecdotal reports of single mortalities (see Section
3.5.1 ). Among these, the most attention-garnering
report was the rediscovery of the black-footed fer-
ret ( Mustela nigripes ), when a farm dog brought in a
freshly killed individual ( Clark, 1987 ; see Gompper,
Preface).
3.3 Exploitative competition
Other than via direct interference competition
and associated indirect effects, dogs can also com-
pete with native carnivores exploitatively for food
( Box 3.1 ). Because domestication has rendered
dogs less ef cient at hunting than wild carnivores
( Clutton-Brock, 1995 ), they are less likely to be ef-
fective exploitative competitors ( Petren and Case,
1996 ; Vanak and Gompper, 2009b ). However, most
populations of dogs are subsidized by humans and
thus can attain numerical superiority, allowing
them to potentially outcompete native carnivores
in acquiring limited resources (as per the abun-
dance-asymmetry hypothesis; Vázquez et al., 2007 ).
Furthermore, the plasticity of dog behavior and the
advantages of food security from human subsidies
can potentially allow dogs to compete with a wide
range of carnivore species, from top-predators such
as wolves, lions, and leopards to mesopredators
such as jackals and foxes as well as nonmammalian
carnivores such as varanid lizards.
Although there are several examples worldwide
of dogs killing often substantial numbers of prey
and being an important cause of mortality for many
species (Ritchie et al., Chapter 2 ), the evidence for
dogs being effective exploitative competitors is not
robust. This is mainly due to the fact that most dog
3.2.3 Dogs as predators of other carnivores
The killing and eating of one carnivore by another is
termed intraguild predation ( Polis et al., 1989 ). Sev-
eral reviews have highlighted the widespread oc-
currence of this interaction ( Donadio and Buskirk,
2006 ; Glen and Dickman, 2005 ; Ritchie and Johnson,
2009 ). However, we draw the distinction between
‘routine’ predation, where the victim is killed main-
ly for consumption, from killing as a form of ex-
treme interference competition, where the victim is
not consumed or only partially consumed (see also
Butler et al., Chapter 5 ). There are numerous exam-
ples of dogs killing other carnivores, both with and
without consumption. For example, in Australia,
dingoes have been recorded killing but not eating
spotted-tailed quolls ( Dasyurus maculatus ) ( Körtner,
2007 ; Körtner and Watson, 2006 ), red foxes, and
feral cats ( Moseby et al., 2012 ), but more generally
they appear to eat or even cache their victims. Other
than quolls, large varanids and snakes have been re-
corded in the diet of dingoes in several studies (e.g.,
Glen et al., 2011 ; Pascoe et al., 2012 ). However, the
frequency of occurrence of other predators in dingo
scats or stomach samples is usually very low (<1%),
and it is seldom clear whether dingoes have killed
or merely scavenged these items. One exceptional
observation is that of Webb ( 1996 ) who saw three
dingoes work together to kill and then consume a
large (1.5–2 m long) lace monitor ( Varanus varius ).
Evidence of killing by dogs comes from almost
everywhere they have been studied, and the list of
mammalian carnivore species that are killed by dogs
is long. The list includes taxa with a wide range of
body sizes from dhole ( Cuon alpinus ) ~18 kg; ( Wil-
liams, 1935 ), to civets ~1–6 kg ( Ashraf et al., 1993 ;
Dahmer, 2002 ), small cats ~5 kg ( Barashkova and
Smelansky, 2011 ) and mustelids ~1–2 kg ( Butler and
du Toit, 2002 ; Maran et al., 2009 ; Ross et al., 2008 ).
In many cases intraguild killing (and not preda-
tion) can be deduced because the carcasses of even
small carnivore species are recovered relatively in-
tact. For example, dogs were the presumed cause
of direct killing of nearly 29% of Indian foxes that
were found dead ( n = 13) ( Vanak, 2008 ). Similarly,
dogs killed 26% of spotted skunks ( Spilogale puto-
rius ) known to have died ( n = 77) during a study
conducted in farmlands in southeast Iowa ( Crabb,
04-Gompper-Chap03.indd 8004-Gompper-Chap03.indd 80 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 81
India, dogs dominated cattle carcasses at a village
dump throughout the day, preventing scavengers
such as golden jackal ( C. aucreus ), vultures, and
striped hyena ( Hyaena hyaena ) from accessing these
resources (Aiyadurai and Jhala, 2006).
A more detailed investigation into the potential
scavenging interactions between dogs and wild car-
nivores was carried out in Zimbabwe (see Section
3.2.2). At experimental carcasses, Butler and du Toit
( 2002 ) found that dogs and vultures were the most
successful species in the scavenging guild, which
also consisted of eight mammalian and nine avian
species. Dogs were the most ef cient scavengers,
consuming up to 60% of the total carcass biomass,
compared to 15% for the vultures. However, the in-
uence of dogs varied along transects. Dogs con-
sumed the majority of carcass biomass 1 km within
the communal land, whereas vultures, leopards,
lions, and spotted hyenas were the principal scav-
engers at carcasses placed 1 km inside the wildlife
reserve.
These patterns suggest that dogs primarily com-
pete with vultures along the interface of communal
lands and wildlife reserves. While most carnivores
will scavenge given the opportunity, vultures have
evolved speci cally as scavengers of mammalian
carrion, and the eagle-like vultures (hooded, Ne-
crosyrtes monachus , white-headed, Trigonoceps oc-
cipitalis , and lappet-faced, Torgos tracheliotus ) will
also scavenge carrion and human refuse ( Houston,
1979 ; Mundy, 1982 ). Dogs appear to out-compete
vultures for four reasons. First, they physically
dominate vultures at carcasses, aggressively ex-
cluding them from feeding due to their larger body
mass (15 kg versus 6.2 kg for lappet-faced vultures,
the largest species recorded scavenging; Houston,
1979 ). Second, because the experimental carcasses
were small (<50 kg, as with the majority of carcasses
found in GCL; Butler and du Toit, 2002 ), dogs were
easily able to defend carcasses against vultures, and
consume them quickly. Third, although vultures are
attracted to carcasses in GCL, they are more sensi-
tive to human disturbance and reluctant to settle
at carcasses near human settlements. The majority
of human activity occurs during the day, the only
time when vultures forage. Fourth, dogs are ac-
tive for most of the 24-hour cycle, enabling them to
populations are heavily dependent on HDF. Vanak
and Gompper ( 2009b ) found that of 21 studies of
dog diet, only 5 (23%) reported subsistence on
mainly wild-caught food whereas in the rest dogs
were either fully or mainly dependent on HDF. Il-
lustratively, all of those  ve studies were of truly
feral dogs (i.e., dogs that were no longer tame or
associated with human settlements and food subsi-
dies). Even studies that have speci cally examined
the potential for exploitative competition between
dogs and sympatric carnivores have found little
evidence. Atickem et al. ( 2010 ) found that dogs in
the Ethiopian highlands subsisted almost entirely
on HDF, and that wild-caught prey such as rodents
constituted a relatively minor part of their diet. The
authors thus concluded that dogs are unlikely to
be competing with endangered Ethiopian wolves.
Similarly, in India, dogs and Indian foxes did not
appear to compete directly for food. This was again
due mainly to the fact that dogs subsisted almost
entirely on HDF such as garbage, human feces, di-
rect feeding, and carrion, whereas foxes subsisted
almost entirely on rodents, fruit, and invertebrates
( Vanak and Gompper, 2009a ). Silva-Rodríguez et al.
( 2010a ) found similar low levels of overlap in diet
between dogs and chilla foxes in southern Chile.
On the other hand, Campos et al. ( 2007 ) reported
high rates of vertebrate consumption by dogs in
suburban areas of Brazil. Although these authors
suggested that dogs could compete with wild car-
nivores, there was some uncertainty in this regard
since there was no assessment of dietary overlap.
3.3.1 Dogs competing with mammalian
and non-mammalian scavengers
Almost all studies that examine the dietary habits of
dogs conclude that an important part of their diet is
obtained from scavenging human- and non-human
generated carcasses of animals. This highly ef cient
facultative scavenging behavior brings dogs into
direct competition with a suite of mammalian and
non-mammalian species that are either specialist or
opportunistic scavengers. Many of the same factors
described previously, such as high population den-
sities, proximity to humans, and 24-hour foraging
ability, allow dogs to dominate carcasses, especially
near human settlements. For instance, in western
04-Gompper-Chap03.indd 8104-Gompper-Chap03.indd 81 14/09/13 9:34 AM14/09/13 9:34 AM
82 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
affect populations within wildlife reserves. The
competitive impact of dogs on the smaller and more
omnivorous brown ( H. brunnea ) and striped hyenas
may potentially be more signi cant, but no research
has been undertaken on this question.
3.3.2 Dogs competing as top-predators
The success of dogs as exploitative competitors
seems to depend to a large extent on their associa-
tion with humans. As mentioned earlier, in the re-
view conducted by Vanak and Gompper ( 2009b ),
only 5 of 21 studies of dog food habits showed a
primary dependence on wild-acquired food. It is
notable that four of these dog populations were ful-
ly independent of humans and acted as wild preda-
tors. Thus, the true competitive ability of dogs with
other carnivores can be assessed in situations where
dogs are wild. Such a situation is best seen on a few
islands (e.g., Kruuk and Snell, 1981 ) and on the con-
tinent of Australia.
Aside from the thylacine and Tasmanian devil,
there is no evidence that any other predatory na-
tive species declined or became extinct in Australia
following the introduction of the dingo, suggesting
that competition between dingoes and extant native
species may not be strong. Although contemporary
studies generally support this interpretation, there
is nonetheless considerable evidence that dingoes
share food, habitat, or shelter resources with many
native species. Hence, the potential for resource
competition exists. Dingoes show some overlap in
diet with spotted-tailed and western quolls ( D. geof-
froii ); all species prey upon small to medium-sized
mammals and birds, although dingoes are capable
of hunting much larger prey than the marsupials,
especially when in packs ( Corbett, 1995 ; Glen and
Dickman, 2008 ; Glen et al., 2011 ). It is also possible
that the marsupial predators consume the remains
of dingo-kills and gain an energetic bene t from
doing so ( Dickman, 1992 ). However, good evidence
for this is scant; large mammals such as sheep have
been identi ed in the diet of the spotted-tailed
quoll, but it is not clear whether these prey had been
killed initially by dingoes or had been scavenged
after death from other causes ( Glen et al., 2011 ).
Large predatory birds such as wedge-tailed ea-
gles ( Aquila audax ) also share prey including rabbits
locate and then consume carrion more ef ciently
than vultures.
Similar evidence also points to dogs out- competing
black vultures ( Coragyps atratus ) in South America.
According to Pavés et al. ( 2008 ) , dogs reduce food
availability for vultures by preying and scaveng-
ing on South American sea lions ( Otaria  avescens ).
Similarly, with the collapse of the vulture popula-
tion in south Asia, several reports suggest that dog
numbers have increased substantially and have re-
placed vultures as the main scavengers ( Prakash
et al., 2003 ).
By being effective scavengers, dogs can poten-
tially out-compete much larger carnivores. For ex-
ample, in Africa lions and leopards also scavenge
opportunistically ( Kruuk, 1972 ; Schaller, 1976 ;
Smith, 1977 ) and considering that both species prey
on dogs ( Butler et al. 2004 , Chapter 5 ), they could
certainly outcompete dogs during direct interfer-
ence confrontations. In spite of this, in Zimbabwe
dogs were the more ef cient scavengers at experi-
mental carcasses owing to their higher numbers,
enabling them to locate and consume larger quan-
tities of carrion. Consequently dogs probably do
successfully compete with lions and leopards in ex-
ploitative terms, but because both felid species are
primarily predators (rather than scavengers), the
impact of dogs is likely to be minimal. There is little
evidence to show that dogs effectively kleptopara-
sitize recent kills of wild carnivores.
Spotted hyenas present a more complex issue.
They are large (up to 86 kg) and highly ef cient
scavengers, and are able to locate and consume tis-
sue and bones from carcasses of all but the largest
species ( Richardson, 1980 ). They often scavenge
refuse around human settlements, for example in
Ethiopia ( Kruuk, 1968 ), Tanzania ( Kruuk, 1972 ),
and South Africa ( Pienaar, 1969 ), but are also high-
ly successful predators in their own right ( Kruuk,
1972 ; Mills, 1990 ). Given that they prey on dogs in
Zimbabwe ( Butler et al., 2004 ), Tanzania ( Kissui,
2008 ), Kenya ( Kolowski and Holekamp, 2006 ) and
Ethiopia ( Atickem et al., 2010 ) (see also Butler et al.,
Chapter 5 ), they are also likely to dominate dogs in
direct competitive encounters involving scaveng-
ing. Due to their higher densities, dogs may po-
tentially reduce food supplies for spotted hyenas
around human settlements, but this is unlikely to
04-Gompper-Chap03.indd 8204-Gompper-Chap03.indd 82 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 83
and procyonids. The spillover of these pathogens
from dogs to wild carnivores and the consequences
of these events for the population dynamics of wild
carnivores have been reviewed extensively ( Cleave-
land et al., 2007 ; Deem et al., 2000 ; Funk et al., 2001 ;
Laurenson et al., 1998 ; Knobel et al., Chapter 6 ).
3.5 Conservation implications
3.5.1 Dog impacts on threatened carnivores
Most of the directed studies on the impacts of dogs
on carnivores have used non-threatened species as
study models (e.g., Silva-Rodríguez et al., 2010a ;
Vanak and Gompper, 2010 ). However, dogs have
been reported to kill a large number of threatened
carnivores (either for consumption or intraguild
killing). To our knowledge there is (mostly an-
ecdotal) evidence of dog-caused mortality for at
least 14 (23%) of 61 species of extant threatened
terrestrial carnivores (Silva, 2012). For example,
in Nepal, dogs are thought to be one of the most
frequent causes of mortality of the red panda ( Ailu-
rus fulgens ) ( Williams, 2004 ; Yonzon and Hunter Jr,
1991 ). In southern India, the death of 10 (of 22 dead)
critically endangered Malabar civet ( Viverra civet-
tina ) was attributable to dogs ( Ashraf et al., 1993 ).
In Estonia, dogs killed 4 of 54 European minks
( Mustela lutreola ) that had been released as part of
a mink restoration program ( Maran et al., 2009 ).
In different areas of the world, local people inform
that dogs have killed threatened carnivores such
as the Andean cat ( Leopardus jacobita ), guigna ( L.
guigna ), Darwin’s fox ( Lycalopex fulvipes ), smooth-
coated otter ( Lutrogale perspicillata ), and southern
river otter ( Lontra provocax ) ( Espinosa-Molina,
2011 ; Hon et al., 2010 ; Lucherini and Merino, 2008 ;
Silva-Rodríguez and Sieving, 2011 ; Soler et al.,
2004 ). Other examples of threatened carnivores
with evidence of killing by dogs include the black-
footed ferret ( Clark, 1987 ), dhole ( Williams, 1935 ),
narrow-striped mongoose ( Hawkins, 2008 ; Wool-
aver et al., 2006 ), marine otter ( L. felina ) ( Pizarro-
Neyra, 2008 ), marbled polecat ( Vormela peregusna )
( Dulamtseren et al., 2009 ), and Cozumel raccoon
( Procyon pygmaeus ) ( McFadden et al., 2010 ). Con-
sidering the relative paucity of studies on free-
ranging dogs, it is very likely that the proportion
( Oryctolagus cuniculus ) and small to medium-sized
macropods with dingoes, as do large reptilian preda-
tors, such as varanid lizards ( Varanus spp.) and salt-
water crocodiles ( Crocodylus porosus ) ( Glen et al., 2010 ;
Parker et al., 2007 ; Pascoe et al., 2012 ; Taylor, 1979 ).
During an irruption of native rodents in central Aus-
tralia, Pavey et al., ( 2008 ) reported a dietary overlap of
88% between dingoes and letter-winged kites ( Elanus
scriptus ), which provides considerable potential for
food-based competition. Dingoes occupy the same
broad range of habitats as native predators ( Fleming
et al., 2001 ) and overlap with some species on a  ne
spatial scale, although a degree of segregation via dif-
ferential use of habitat components such as trees is
usually still apparent ( Glen et al., 2011 ). Dingoes also
overlap broadly in their temporal activity with mar-
supial, some avian, and reptilian predators. However,
until simultaneous monitoring of the dingo and native
predators is attempted via remote tracking or camera
trapping, it will not be possible to quantify patterns of
temporal overlap or separation more precisely.
3.4 Apparent competition
The occurrence of apparent competition, wherein
patterns of interaction that seem to be the result of
competition are actually driven by the distribution
and effects of a third species ( Box 3.1 ), has not been
robustly demonstrated among carnivores. How-
ever, apparent competition between carnivores as a
result of shared parasites may be far more common.
Several important microparasites of wild carnivores
are also enzootic in dog populations and may be
readily transmitted to native carnivores (Knobel et
al., Chapter 6 ). For instance, almost all canid spe-
cies are susceptible to infections from rabies virus,
canine distemper virus (CDV), and canine parvo-
virus (CPV), and in some cases these three viruses
are primary drivers of carnivore population dy-
namics ( Cleaveland et al., 2007 ). This potential for
disease-mediated changes in population densities
of carnivores that are sympatric with dogs has been
suggested as a form of apparent competition ( Vanak
and Gompper, 2009b ). Prominent examples of the
role of dogs as reservoirs of pathogens that signi -
cantly impact wild carnivore populations come from
species in several carnivore families including can-
ids, felids, hyaenids, phocids, mustelids, viverrids,
04-Gompper-Chap03.indd 8304-Gompper-Chap03.indd 83 14/09/13 9:34 AM14/09/13 9:34 AM
84 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
of doing this is to use domestic dogs as livestock
guardians, to keep dingoes away from rangeland
enterprises ( Box 3.2 ; van Bommel and Johnson,
2012 ).
Although dogs can help reduce human–
carnivore con ict, they can also intensify it. Just
as for other carnivores, dogs prey on livestock,
but distinguishing between losses caused by
dogs and those by other carnivores is a dif cult
task. Despite the dif culty involved in the cor-
rect determination of cause of death, carnivores
such as wolves are often blamed for losses that
could have been caused by dogs ( Cozza et al.,
1996 ). For example, in northern Spain, wolves
were blamed for 94% of the attacks on domestic
animals ( Cozza et al., 1996 ). Paradoxically, in the
same area and during the same time period, dogs
were more abundant than wolves. Furthermore,
sheep represented 36% of prey items found in dog
scats versus only 3% in wolf scats, thus suggest-
ing that the actual impacts of wolves may have
been overestimated ( Echegaray and Vilà, 2010 ).
These identi cation problems are a major issue
for carnivore conservation, because they may
lead to unjusti ed persecution of carnivores, and
may increase the cost of compensation strategies
to reduce human–carnivore con icts ( Cozza et al.,
1996 ; Echegaray and Vilà, 2010 ; Sundqvist et al.,
2008 ). Another situation where high densities of
dogs can inadvertently trigger human–carnivore
con ict is when they are prey for larger carni-
vores. As described by Butler et al. (Chapter 5),
dogs are killed and often consumed by several
species of large carnivores. In cases where the dog
is a beloved pet or a prized animal, this causes re-
sentment towards the presence of large carnivores
and can be a major impetus for predator removal
programs. Thus, it appears that dogs may be ben-
e cial in mitigating human– carnivore con ict,
but only in a narrow set of conditions where they
are fully under human supervision.
3.5.3 Dogs as a large-scale edge effect
A body of evidence is starting to accumulate on the
existence of large-scale edge effects as a function of
human-driven activities ( Laurance, 2000 ). Contrary
to previous evidence, edge effects are not con ned
of species of conservation concern killed by dogs is
much higher.
From the information available, the picture that
emerges is that although the most compelling evi-
dence supporting the impacts of dogs come from
non-threatened carnivores such as the Indian fox
( Vanak and Gompper, 2010 ), the same kind of prob-
lem may be affecting species that are critically en-
dangered, such as the Malabar civet, the Cozumel
raccoon, or Darwin’s fox. Research directed at un-
derstanding the effects of dogs on both the distri-
bution and population dynamics of endangered
carnivores is urgently needed, but in the interim,
the information available on common species could,
under the precautionary principle, inform decision-
making (Silva, 2012).
3.5.2 Dogs and human–carnivore confl ict
Under certain conditions, the presence of dogs
may be beneficial for conservation. Human–
carnivore conflicts are one of the main threats for
carnivore conservation worldwide ( Inskip and
Zimmermann, 2009 ; Treves and Karanth, 2003 )
because the economic losses from livestock dep-
redation lead to intolerance of wild carnivores.
Livestock-guarding dogs are most commonly
used to mitigate this conflict (VerCauteren et al.,
Chapter 9 ). In such cases, the function of dogs is
to keep native carnivores away from livestock.
Hence, the negative correlation between the dis-
tribution of carnivores and dogs in many studies
( Espartosa, 2009 ; Silva-Rodríguez et al., 2010a ;
Vanak and Gompper, 2010 ) may ultimately help
in reducing carnivore mortality associated with
human–wildlife conflict, and therefore benefit
conservation in the long run ( González et al.,
2012 ).
In Australia, where the dingo is the ‘top-dog,’
much evidence has emerged in recent studies that
this carnivore provides a net bene t to native wild-
life by suppressing the impacts of the red fox and
perhaps also the feral cat ( Glen and Dickman, 2005 ;
Letnic et al., 2012 ). However, as dingoes also cause
livestock losses in rangeland areas, their manage-
ment for conservation needs to be balanced against
mitigating their effects in production landscapes.
Intriguingly, one of the most promising means
04-Gompper-Chap03.indd 8404-Gompper-Chap03.indd 84 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 85
problematic for conservation, particularly when
dogs occur in and around protected areas (e.g.,
Lacerda et al., 2009 ; Srbek-Araujo and Chiarello,
2008 ; Torres et al., 1996 ; Vanak and Gompper, 2010 ).
However, not all dogs are likely to be equal in their
effects on carnivores. As postulated by Vanak and
Gompper ( 2009 b), the competitive ability of dogs is
likely to be in uenced by their population size and
ranging behavior. The larger the population of dogs,
and the wider ranging their behavior, the more like-
ly they are to either directly or indirectly in uence
other carnivore species. Thus, to negate the possi-
ble deleterious effects of dogs on carnivores, it is
necessary to not only reduce population size, but
also restrain free-ranging behavior. Priority should
be given to male dogs, since they have larger home
ranges than females (Butler et al., Chapter 5 ) and
travel greater distances when rabid (Knobel et al.,
Chapter 6 ), and therefore may be likely to interact
more frequently with wild carnivores. Most of the
emphasis on free-ranging dog populations world-
wide has thus far been on reducing the potential for
zoonotic diseases, in particular rabies. Even in cases
where dogs have been implicated as the source of
pathogens causing widespread mortality in carni-
vores, the emphasis has been on reducing the trans-
mission potential via vaccination (e.g., Cleaveland
et al., 2007 ), rather than on population control or
restricting ranging-behavior.
Mitigating the risk of pathogen transmission does
not, however, reduce the possibility of other kinds
of dog–wildlife interactions ( Vanak and Gompper,
2010 ). As discussed above, these depend on popula-
tion size and ranging behavior. Therefore, control-
ling dog–wildlife interactions, particularly in the
vicinity of conservation areas, must involve a multi-
pronged approach. Pathogen transmission risk can
be mitigated through vaccination, and the biotic po-
tential of the population can be reduced via lethal
control and sterilization. In areas of conservation
concern, control measures must also include the re-
moval of un-owned dogs, restriction of free-ranging
activity (particularly among males), improvement
of feeding, and a strong emphasis on responsible
dog ownership ( Vanak and Gompper, 2010 ; Silva-
Rodríguez and Sieving, 2011 ). These management
approaches must be implemented in a sustained
and integrated manner for a long-term solution.
to just <150 m from a habitat edge, but can poten-
tially extend up to 5 km from a disturbance edge
( Laurance, 2000 ; Murcia, 1995 ). As shown above
(Section 3.2.2), in Zimbabwe dogs were found 3
km within a wildlife reserve independently of their
owners, and 6 km with people, and clearly modi ed
the scavenging guild up to 1 km within the wildlife
reserve. This study also demonstrated that dogs
were the most common carnivore along the wildlife
reserve boundary, occurring with a frequency six-
fold greater than the most common wild carnivore,
the leopard. Fragmentation and the loss of habitat,
along with increasing human populations, have re-
sulted in an ever-increasing proportion of natural
areas subject to anthropogenic edge effects. Several
recent studies have now proposed that domestic
carnivores such as dogs and cats can constitute a
large-scale edge effect ( Torres and Prado, 2011 ).
Apart from these directed studies, a number
of general faunal surveys across the world have
noted that dogs are among the most commonly
detected carnivore species ( Table 3.2 ) in several
natural areas with close proximity to human habi-
tation. Dogs may be detected as far as 10 km from
the nearest human habitation (and much further
away in Australia), and thus the likely impacts
they can have are not necessarily con ned to short
distances from human settlements. For most of the
surveys and studies listed in Table 3.2 , examina-
tion of the effects of dogs on other species was not
their primary goal. Thus, the information gathered
from across these studies is heterogeneous with
regard to detail, and is intended to be purely il-
lustrative. What emerges is that dogs are quite
commonly detected in several surveys. Given the
scope for negative interactions between dogs and
native species discussed above and elsewhere in
this book, there appears to be a legitimate cause
for concern. There is thus not only a need for more
directed research on the extent to which dogs can
be viewed as an edge effect, but also for general
surveys to explicitly report the presence of domes-
tic carnivore species.
3.5.4 Dog management for conservation
The competitive dynamics between dogs and sym-
patric carnivores described above can clearly be
04-Gompper-Chap03.indd 8504-Gompper-Chap03.indd 85 14/09/13 9:34 AM14/09/13 9:34 AM
86 FREE-RANGING DOGS AND WILDLIFE CONSERVATION
Table 3.2 Summary of a sample of surveys that recorded dogs in natural habitats as an example of a large-scale edge effect. We have excluded
studies of dingoes from Australia as they are naturalized predators in the system and may not be considered as an edge effect.
Region Habitat Detection method Frequency of detection
relative to other carnivores
Distance from human
habitation (km)
Reference
Europe
Spain Agricultural matrix Transects (spotlight) Red fox > dog > cat > others Variable Sobrino et al. 2009
Spain Woodland Camera traps Cat > dog> fox > others < 0.4 Fandos et al. 2012
Portugal Mediterranean
farmlands
Sign detection in
transects
Red fox > dog > Egyptian
mongoose > Eurasian badger
> cat > Eurasian otter >
weasel ~ European polecat
~ stone marten > genet
Variable Pita et al. 2009
Asia
Taiwan Rainforest and
secondary forest
Camera traps Dogs detected, frequency not
reported
2–10 Chen et al. 2010
Malaysia Rainforest Transects and
camera traps
Dogs, jungle cats, and common
palm civet frequently detected
<1 Sanei and Zakaria
2011
India Grassland and
plantation
Camera traps and
track plots
Indian fox > dog > others 1–3 Vanak and
Gompper 2007
North America
USA California sage scrub,
annual grassland,
chaparral, and oak
woodland
Camera traps Coyote > bobcat > dog >
striped skunk > raccoon > gray
fox > puma > others
Dogs associated with
urban percentage but not
with distance to edge
Ordeñana et al.
2010
USA Deciduous/coniferous
forests
Camera traps, Scent
stations
Albany Pine Bush Preserve
(APB): Cat > coyote ~ raccoon
> dog > others. Adirondack
sites: dogs not detected.
APB: Close to suburban
and urban development
Gompper et al.
2006
South America
Brazil Atlantic forest Camera traps Dog > ocelot > crab-eating
raccoon > puma > coati >
crab-eating fox > others
0–1.1 from forest edge.
A dog detected 0.9 from
residence.
Srbek-Araujo and
Chiarello 2008
Brazil Cerrado Track stations Maned wolf > dog >
crab-eating raccoon >
crab-eating fox > coati
~ puma ~ tayra
Associated to edge of
National Park (< 1)
Lacerda et al.
2009
Chile Temperate rainforest Camera traps and
track stations
Dog > guigna > Darwin’s fox
> Southern river otter > others
Variable Farías and Jaksic
2011
Chile Temperate rainforest Camera traps Dog > skunk > guigna
~ puma ~ chilla fox
Variable Silva-Rodriguez
et al. 2010b
Africa
Zimbabwe Woodland savannah Transects (tracks on
sand, experimental
carcass observation,
wildlife monitoring
transects)
Dog > leopard > side-striped
jackal > lion > spotted hyena
3 (independent of
humans), 6 (with
humans)
Butler and du Toit,
2002 ; Butler et al.
2004
04-Gompper-Chap03.indd 8604-Gompper-Chap03.indd 86 14/09/13 9:34 AM14/09/13 9:34 AM
COMPETITION BETWEEN DOGS AND SYMPATRIC CARNIVORES 87
behaviour at ungulate carcasses . Animal Behaviour , 75 ,
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Barashkova , A. and Smelansky , I. ( 2011 ). Pallas’s cat in the
Altai Republic, Russia . Cat News , 54 , 4 – 7 .
Barcala , O. ( 2009 ). Invasive stray and feral dogs limit fossa
( Cryptoprocta ferox ) populations in Ankarafantsika Na-
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tor distribution around water points in arid Australia .
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fects of predator control on behaviour of an apex preda-
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Butler , J.R.A. ( 1998 ). The ecology of domestic dogs Canis
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3.6 Further research
This review highlights the general lack of studies
on the competitive dynamics between dogs and
sympatric carnivores, both in terms of geographic
representativeness as well as the range of species
that are affected. Indeed, barely a handful of studies
from Asia, Africa, and South America have directly
examined these competitive dynamics. Even when
direct aggression and intraguild predation can be
detected among competing species, the subtler in-
direct effects of exploitation and apparent compe-
tition may sometimes escape notice. The various
competitive roles of dogs are still largely unknown.
As discussed in this review, dogs may have large-
scale effects on native carnivores despite not com-
peting strongly for food. Hence, it is clear that there
are major gaps in directed research on the effects of
dogs on sympatric carnivores worldwide, but par-
ticularly in Asia, Europe, and Africa, which togeth-
er have the largest populations of dogs as well as a
wide array of native carnivore species.
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