House cats as predators in the Australian environment: impacts and management

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House cats as predators in the Australian environment: impacts House cats as predators in the Australian environment: impacts

and management and management

Christopher R. Dickman

University of Sydney, New South Wales, Australia

, [email protected]

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Human–Wildlife Conﬂ icts 3(1):41–48, Spring 2009

House cats as predators in the Australian

environment: impacts and management

CHRISTOPHER R. DICKMAN, Institute of Wildlife Research, School of Biological Sciences, Uni-

versity of Sydney, New South Wales 2006, Australia [email protected]

Abstract: This paper provides an overview of the predatory activities of the house cat (Felis

catus) in Australia, focusing principally on the interactions of domestic and stray cats with

native species of prey. Like their free-living, or feral, counterparts, domestic cats take a broad

range of prey, with small mammals, birds, and human-derived foods forming the bulk of the

diet. Domestic and stray cats have contributed to declines of suburban populations of eastern

barred bandicoots (Perameles gunnii) and superb lyrebirds (Menura novaehollandiae) in

Victoria, Australia. The effects of cats on prey communities remain speculative. In Sydney,

artiﬁ cial nests placed in trees in forest remnants suffered less predation where cat activity

was high rather than where it was low, indicating that cats beneﬁ cially reduced damage by

introduced rats and other nest predators. However, high cat activity was associated with

reduced bird diversity. Legislation to encourage responsible cat ownership has been passed

in Australia; it should have positive outcomes for both wildlife conservation and cat welfare.

Key words: Australia, Felis catus, house cat, human–wildlife conﬂ icts

Over the last 20 years, there has been a

surge of interest in the introduced house cat

(Felis catus) in Australia and, in particular, the

impact of the cat on native Australian fauna.

Some studies have suggested that cats kill

millions of native vertebrates each year (Paton

1990, 1991; Trueman 1991), and that certain

species are represented disproportionately

in the kill (Seebeck et al. 1991, Dowling et al.

1994). Predation from cats appears to have been

a major contributor to declining populations

of some threatened native species, such as the

eastern barred bandicoot (Perameles gunnii)

in Victoria and the rufous hare-wallaby

(Lagorchestes hirsutus) in the Northern Territory

(Du y 1994, Gibson et al. 1994). Losses of

such species have led to a empts to extirpate

cats from local (<10,000 ha) or even regional

(>10,000 ha) areas (Algar et al. 2002, Short et

al. 2002, Short and Turner 2005), and many

a empts have been made to design cat-speciﬁ c

traps, lures, or toxins (Algar and Burrows 2004,

Wark 2004).

Other studies have argued, conversely, that

the kill-rates of cats are lower than o en is

believed, especially in urban environments

(Reark 1994), and that introduced vertebrates

usually form the major part of their diet (Barra

1997). Cats o en are perceived to be beneﬁ cial

as controllers of vermin on farms and rural

properties (Ward 1994), and possibly to have

positive eﬀ ects on some native species by

suppressing populations of introduced Ra us

spp. (Tidemann et al. 1994). In Australia, where

cats were introduced in large numbers in the

eighteenth and nineteenth centuries (Abbo

2002), negative impacts have been highlighted

most o en. The strong public aﬀ ection for cats

(e.g., Murray and Penridge 1997, Grayson et al.

2002) and limited empirical evidence of their

actual impacts have hampered a empts to

manage them eﬀ ectively.

Studies of cat impact o en have drawn a

distinction between 2 kinds of cats. On the

one hand, domestic cats have been viewed as

pet or house cats that live in close connection

with a household where all their ecological

requirements are intentionally provided by

humans (Moodie 1995). Such cats do not rely on

hunting for food, but they may still impact on

native fauna by their predatory activities. On the

other hand, feral cats are free-living; they have

minimal or no reliance on humans, and survive

and reproduce in self-perpetuating populations

House cat. (Photo courtesy P. German)

Human–Wildlife Conﬂ icts 3(1)

(Moodie 1995). Individual cats may sometimes

move between these 2 extremes, occupying the

category of stray if there is partial dependency

on humans for the provision of resource

requirements.

In the present paper, I discuss the predatory

activities of cats in Australia and review the

interactions of cats with populations and

communities of native prey species. I address

5 questions:

What do cats eat?1.

Are cats specialist or generalist pred-2.

ators?

What are the eﬀ ects of cat predation on 3.

prey populations?

What are the eﬀ ects of cat predation on 4.

prey communities?

What management protocols can be im-5.

plemented to mitigate the impacts of

cats?

I focus primarily on predation by domestic

and stray cats because the impacts of feral cats

on native fauna have been much studied and

reviewed (e.g., Dickman 1996a, b; Risbey et al.

1999, 2000) and are currently the subject of a

national threat abatement plan in Australia

(Environment Australia 1999). In addition, there

is emerging evidence that owned and stray cats

o en disperse into the natural environment

and help to sustain feral populations (Denny

et al. 2002, Hutchings 2003), and so contribute

more to impacts on native fauna than has

been realized hitherto. There has been limited

documentation of the eﬀ ects of domestic

and stray cats on native fauna (e.g., Po er

1991, Siepen and Owens 1993, Paxton 1994). I

review this information here, but also present

new information from my own studies where

possible.

What do cats eat in Australia?

While cats take a broad range of prey,

small mammals and birds o en feature most

prominently in their diet. In a summary of 22

studies of the diet of feral cats from 20 localities

in mainland Australia, Dickman (1996a)

found that introduced rabbits (Oryctolagus

cuniculus) and house mice (Mus musculus)

are major dietary items in semi-arid and arid

habitats, whereas marsupials predominate

in temperate forest habitats. In both forest

and suburban habitats, the common ringtail

possum (Pseudocheirus peregrinus) is depredated

frequently. On islands of the Australia-Paciﬁ c

region, birds o en predominate in the diet of

cats (Fitzgerald 1990, Dickman 1996a), although

invertebrates also are a prominent part of their

diet (Fitzgerald and Veitch 1985, Hayde 1992).

Studies published since 1996 support the view

that cats take a broad range of prey (Barra

1997, Meek 1998, Murphy et al. 2004), including

carrion, under certain conditions (Paltridge et

al. 1997, Molsher et al. 1999).

Studies of cat diet in suburban and temperate

forest habitats of the Sydney Basin, New South

Wales, exemplify the range of prey taken by

domestic and stray cats (Table 1). In the most

suburban situation, at Cooper Park, Sydney,

native vertebrates (small scincid lizards, birds,

and the common brushtail possum [Trichosurus

vulpecula]) together formed as li le as 8–17% of

the diet of cats by volume. Most of the diet was

derived from human-provided sources of food.

In contrast, at suburban North Head, Sydney,

and in temperate forest at Olney and Kurin-

gai on the city’s northern fringe, mammals

predominated in the diets, with 37 to 60% of

the total volume being derived from native

species (Table 1). Dietary diﬀ erences among

habitats probably reﬂ ect diﬀ erences in prey

availability. The common brushtail possum is

the only terrestrial native mammal occurring in

Cooper Park and surrounding areas, whereas

native mammals predominate at the other sites

(Ma hews et al. 1999). Rabbits are likewise

abundant at North Head, where they form 21%

of the diet, by volume, of cats, but scarce at the

other sites, where they form only 1 to 6% (Table

1). These results support previous ﬁ ndings

(Fitzgerald 1988) that cats prey mostly on small

vertebrates, especially mammals and birds.

These results also are consistent with studies

elsewhere in Australia (Wallis et al. 1996) and

Europe (Goldschmidt-Rothschild and Lüps

1976, Borkenhagen 1979) that demonstrate

large diﬀ erences in the representation of prey

species between suburban and less disturbed

natural habitats.

Are cats specialist or generalist

predators?

The predominance of small mammals and

43House cats • Dickman

birds in the diet of cats has led to the frequent

assumption that these taxa are preyed upon

selectively. However, dietary selectivity is more

reliably indicated if a predator can be shown

to take certain prey over others that are also

available. Such selectivity may be demonstrated

at the level of the individual predator or that

of the population. In a particularly instructive

study of population-level selectivity, Childs

(1986) showed that cats in urban Baltimore,

Maryland, USA, took small brown rats (Ra us

norvegicus) weighing <200 g, while most (91%)

of the rats available in the population weighed

>200 g. In other studies, Barra (1997) showed

that cats in suburban Canberra, Australia,

preferred house sparrows (Passer domesticus) and

blackbirds (Turdus merula) to 9 other common

species of birds, but avoided starlings (Sturnus

vulgaris), possibly because starlings nest and

roost high in trees and feed on the ground in

ﬂ ocks that can more readily detect approaching

predators. Molsher et al. (1999) demonstrated

that rabbits were being depredated selectively

at Lake Burrendong, near Wellington, New

South Wales, even a er a 90% decline in rabbits’

abundance due to rabbit calicivirus disease. At

the individual level, cats have sometimes been

shown to selectively take certain prey species

and to adopt specialized hunting methods to

obtain them. O en these are species of small

mammals or birds (Bradshaw 1992), but large

animals such as the rufous hare-wallaby

(Lagorchestes hirsutus; Gibson et al. 1994) and

unusual prey, such as bats (Churcher and

Lawton 1989) and grasshoppers (Hochstrasser

1970), also have been targeted. Observations of

hunting by domestic and stray cats near human

se lement at 2 locations have revealed clear

specializations among individuals. On Ro nest

Island, Western Australia, 3 of 5 cats appeared to

be accomplished mousers, catching every house

mouse (Mus musculus) that they hunted, but

achieving only a 50% catch rate for either birds

or lizards. These cats adopted a sit-and-wait

strategy (Turner and Meister 1988), pouncing

on mice from behind cover under conditions of

semi-light or darkness. In the same population,

by contrast, 1 cat achieved high rates of capture

success on lizards and another cat on birds.

Both foraged diurnally, the ﬁ rst pursuing lizard

prey actively in sparse coastal heath vegetation,

the second pouncing on birds from dense cover.

Similar selectivity was observed by 1 cat at

North Head that specialized on rabbits.

Table 1. Diets of house cats (Felis catus) in suburban and temperate forest habitats of the Sydney Ba-

sin, New South Wales, shown as percentage volume of occurrence of food categories.

Suburban habitat Forest habitat

Food category Cooper Park

= 37)

North Head

(n = 24)

Olney State Forest

(n = 12)

Kuring-gai National

Park (n = 28)

Rabbit 1.3 21.5 1.3 6.3

House mouse 1.3 1.0 1.9

Other rodent

14.1 22.6 15.8 18.8

Bat 0.7

Marsupial 4.7 37.2 32.8 40.8

Bird 8.8 8.1 27.6 12.4

Reptile 3.6 3.9 4.2 6.0

Invertebrate 5.5 3.7 3.0 2.7

Scavenge 5.0 7.6 1.9

Other

57.0 1.7 6.6 8.5

Sample sizes, n, represent pooled results from analyses of feces at all sites and stomach contents at all

sites except Cooper Park.

In the suburban habitats, all “other rodents” ingested were introduced black rats (Ra us ra us),

whereas in the forest habitats all “other rodents” taken were probably native bush rats (R. fuscipes).

The category “other” represents cat fur, plant material, nonorganic items, unidentiﬁ ed materials, and

foods likely to be of human origin, such as ﬁ sh, bread, or commercial pet food. Full methodological

details are given in Dickman (1996a).

Human–Wildlife Conﬂ icts 3(1)

What are the effects of cat

predation on prey populations?

The population-level impacts of feral cats

on native fauna have been much discussed in

Australia (Dickman 1996a, b) and elsewhere

(King 1984, Fitzgerald 1988, 1990). Surprisingly,

the impacts of domestic and stray cats have

remained poorly studied in Australia, with

only 2 well-documented studies.

The ﬁ rst study concerned the eastern barred

bandicoot (Perameles gunnii), a smal, rabbit-size

marsupial, at the town of Hamilton, Victoria.

Formerly widespread in southwestern Victoria

and southeastern South Australia, this bandicoot

had become restricted to the Hamilton area by

the 1970s. During 1982 to 1983, the population

comprised about 1,750 animals (Moon 1984),

but had fallen to just 150 to 300 animals by

1989 (Lacy and Clark 1990). The precipitous

population decline appeared to be driven by

a high rate of mortality, especially of juveniles,

with >42% of juvenile deaths being caused by

cat predation (Du y 1994).

The second case study concerned the superb

lyrebird (Menura novaehollandiae) in Sherbrooke

Forest, Victoria. The lyrebird population was

about 130 animals in the 1960s, but had fallen

to only 60 by 1988 (Bradley and Bradley 1990).

Predation by cats, foxes (Vulpes vulpes) and

domestic dogs (Canis familiaris) was identiﬁ ed

as the major cause of the decline, with cats

probably accounting for disproportionate

mortality of young birds (Larkin 1989; H.

Bradley, personal communication).

What are the effects of cat

predation on prey communities?

Given the relatively small amount of

information on the eﬀ ects of cats on individual

native species, it is not surprising that

understanding of cat impacts on prey com-

munities is meager. There is considerable

speculation that cats may have indirect but

deleterious eﬀ ects on plant communities by

reducing the abundance of avian pollinators,

or by depleting rat kangaroos (Be ongia

spp. and Potorous spp.) and other vectors

of mycorrhizal fungi (Dickman 1996a). In

contrast, circumstantial evidence suggests that

cats may facilitate denser populations of forest

birds by suppressing the numbers of predatory

rats (Fitzgerald and Karl 1979, Ebenhard 1988,

Tidemann et al. 1994).

One observational study on birds in Sydney

illustrates the diﬃ culty of disentangling the

positive and negative eﬀ ects of cats at the

community level. In the ﬁ rst part of this study,

in which I participated, Ma hews et al. (1999)

investigated the intensity of predation on artiﬁ -

cial bird eggs and nests placed in trees in 24 forest

patches throughout the Sydney metropolitan

region. The predation rate, calculated as the

percentage of nests a acked by predators,

ranged from 45 to 100%. Avian predators were

detected in all patches; black rats (Ra us ra us)

a acked nests in 10 areas, and common ringtail

possums and brown antechinus (Antechinus

stuartii) damaged eggs in 2 areas each. In the

second part of the study, I walked along foot

tracks through each remnant and counted the

numbers of cat feces encountered to obtain a

rough index of cat activity. A plot of cat activity,

expressed as feces-per-km of track, against

nest predation rate revealed a strong negative

correlation (Figure 1). As cats did not damage

any nests in the 24 sites, the reduced levels of

nest predation associated with high cat activity

presumably reﬂ ect suppressive eﬀ ects of cats

on nest predators. Indeed, examination of the

contents of the collected cat feces indicated that

all of the nest predators noted above themselves

fell victim to cats (Dickman, unpublished data),

supporting the presumption that suppression

is likely to have occurred. On their own, these

ﬁ ndings indicate that domestic and stray cats

may beneﬁ t tree-nesting birds in remnant forest

patches in Sydney by reducing rates of nest

predation.

In the ﬁ nal part of this study, the richness

of all native species of birds was sampled in

the same 24 forest patches by scoring species

observed along foot tracks and plo ing them

against cat activity (Figure 2). The strong

negative relationship suggests that cats reduce

the total numbers of bird species that occur

in forest remnants. Species that were absent

from sites with high cat activity, but present

elsewhere, included wrens (Malurus cyaneus),

thornbills (Acanthiza chrysorrhoa, A. pusilla),

wagtails (Rhipidura leucophrys), and other vul-

nerable small species that feed or nest close to

the ground. These observations support the

notion that cats depleted the avian community,

presumably by direct predation. Conﬁ rmation

awaits appropriate experimental studies.

45House cats • Dickman

What management protocols

can be implemented to mitigate

the impacts of cats?

Taken together, the ﬁ ndings of the above

studies provide some evidence that domestic

and stray cats impact negatively on some

native species. If these cats help to establish

and maintain feral cat

populations in less disturbed

habitats outside conurbations

(Denny et al. 2002, Say et al.

2003), their impacts may be

more subtle, but also more

pervasive, than realized.

Research on this issue is

continuing (Denny 2005). In

built-up urban and suburban

environments, it is likely

that any direct impact of

cats on native fauna will

be secondary to the more

dramatic eﬀ ects of loss

and modiﬁ cation of native

vegetation by the suburbs

themselves. However, in less

disturbed areas adjoining

reserves, national parks, or in

remnants of native vegetation

adjoining new residential

developments, predation by

domestic and stray cats on

native species may be quite

damaging (Barra 1997).

In response to community

perceptions about maraud-

ing cats, city councils and

governments in all Austral-

ian states and territories have

debated or passed by-laws

to encourage responsible

cat ownership (Department

of Local Government 1994,

Seebeck and Clunie 1998).

Many municipalities also

provide information packs

to increase the awareness of

owners about cat–wildlife

interactions. By-laws vary

greatly from council to

council, but most include

provisions for registration

of pet cats, incentives for

sterilization, nigh ime curfews, and stipulations

for a maximum number of cats per property;

some also allow for removal of unowned cats

from parks and other areas of sensitive habitat.

Community surveys generally indicate

strong support for legislation that promotes

informed cat ownership, but weaker support for

P < 0.001

Figure 1. Relationship between house cat (Felis catus) activity and

nest predation rate in 24 remnant patches of forest in Sydney, New

South Wales, Australia. Cat activity is expressed as an index based

on numbers of cat feces found per km of foot tracks within patches.

Nest predation rate is expressed as the percentage of artiﬁ cial nests

depredated per patch. Based on Mattews et al. (1999) and Dickman

(unpublished data).

Figure 2. Relationship between house cat (Felis catus) activity and bird

species richness in 24 remnant patches of forest in Sydney, New South

Wales. Cat activity is expressed as an index based on numbers of cat

feces found per km of foot tracks within patches. Bird species richness

is expressed as the number of bird species heard or observed along

foot tracks within patches pooled over 2 to 4 searches per patch.

r = 0.63, P < 0.001

Human–Wildlife Conﬂ icts 3(1)

proposals that restrict ownership or create cat-

free zones (e.g., Grayson et al. 2002). Eﬀ ective

provisions should have twofold beneﬁ ts. First,

they should reduce the depletion of native

wildlife in se led areas. Secondly, they should

improve cat welfare by reducing the numbers

of dumped, unwanted cats, and by reuniting

lost pets with their owners. Despite the broad

community support for education and control

of cats, and the plethora of by-laws that has

been passed in recent years, there has been no

evaluation to date of the eﬀ ectiveness of any

existing programs. Such a review should be

carried out as a ma er of priority.

Acknowledgments

I thank C. McKechnie for assistance with

much of the work described here, the Australian

Research Council for funding, M. Crowther for

comments on the manuscript, and T. English

and S. Roy for encouragement to present this

contribution.

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CHRISTOPHER R. DICKMAN is a professor

of ecology at the School of Biological Sciences and

director of the Institute of Wildlife Research, both at

the University of Sydney, Australia. He is a fellow of

the Royal Zoological Society of New South Wales.

In his research, he investigates factors that inﬂ u-

ence the distribution and abundance of terrestrial

vertebrates. He is curious to know the causes of the

intriguing patterns of vertebrate distributions in the

Australian fauna and is concerned that many spe-

cies have declined or become extinct with the ad-

vent of European settlement. For the last 20 years,

he has studied the exceptionally rich communities

of small mammals and lizards of arid Australia. This

research contributes to theoretical debate about the

importance of biotic and physical processes in shap-

ing population and species dynamics and to practi-

cal conservation gains. He was chair of the NSW

Government Scientiﬁ c Committee for seven years,

sits on several scientiﬁ c advisory bodies, such as

the Science Advisory Committee for Earthwatch,

and is a governor for WWF-Australia. He has served

on editorial advisory boards for several journals, and

is currently an associate editor of the Journal of Ap-

plied Ecology and the Open Ecology Journal. (Photo

courtesy A. Greenville)