Blackwell Publishing LtdOxford, UKMAMMammal Review0305-1838Blackwell Publishing Ltd, 2005? 2005353&4302312review articlePredation by domestic catsP. J.

Baker
et al.

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Mammal Rev. 2005, Volume 35, No. 3&4, 302–312. Printed in Singapore.

Impact of predation by domestic cats Felis catus in an urban area

PHILIP J. BAKER , AMY J. BENTLEY , RACHEL J. ANSELL and

STEPHEN HARRIS
School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK

ABSTRACT
1. As companion animals, domestic cats Felis catus can attain very high densities, and have
the potential to exert detrimental effects on prey species. Yet, there is a paucity of information
on the impact of cat predation in urban areas, where most cats are likely to be present.
2. We quantified the minimum number of animals killed annually by cats in a 4.2-km2 area
of Bristol, UK, by asking owners to record prey animals returned home by their pets.
The potential impact of cat predation on prey species was estimated by comparing the
number of animals killed with published estimates of prey density and annual productivity.
3. Predator density was 229 cats/km2.
4. Five mammal, 10 bird and one amphibian prey species were recorded. Mean predation
rate was 21 prey/cat/annum. The most commonly recorded prey species was the wood mouse
Apodemus sylvaticus.
5. Predation on birds was greatest in spring and summer, and probably reflected the killing
of juvenile individuals. For three prey species (house sparrow Passer domesticus, dunnock
Prunella modularis, robin Erithacus rubecula), estimated predation rates were high relative to
annual productivity, such that predation by cats may have created a dispersal sink for juveniles
from more productive neighbouring areas. The impact of cats on these species therefore
warrants further investigation.

Keywords: conservation, feral cat, house cat, hyperpredation, urban ecology

INTRODUCTION
Hyperpredation is the process whereby one prey species exerts an indirect effect on a second
species by causing an increase in the abundance of a common shared predator. For verte-
brates, this process has generally been considered in the context of the detrimental impact of
introduced wild animals on endemic species (e.g. Courchamp, Langlais & Sugihara, 1999,
2000; Roemer et al., 2001). However, similar effects may arise where the predator is a domestic
or companion animal of humans, with human provisioning of food analogous to the role of
the primary prey species (see also Kristan & Boarman, 2003). Furthermore, because of their
reliance on artificial food sources, predator abundance may substantially surpass the carrying
capacity of the environment, such that the impact of predation may be severe, even though
artificial food may reduce individual predation rates. The most widespread example of such
a system is that of the domestic cat Felis catus.
Cat populations exhibit varying degrees of dependence on humans. Truly, feral populations
live completely independently of humans, whereas urban colonies, farm colonies and stray
cats are generally reliant in part on people for food. However, in many developed countries,

Correspondence: P. J. Baker. E-mail: [email protected]

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Predation by domestic cats 303

the largest fraction probably comprises free-ranging pet or house cats that are regularly fed
by their owners. In the UK, there are estimated to be around 813 000 feral (Harris et al.,
1995) and eight million pet cats (Cats Protection League, 1993).
Individual pet cats vary considerably in the degree to which they predate wild animals,
with most probably taking few or no prey (Churcher & Lawton, 1987; Barratt, 1997, 1998;
Robertson, 1998; Ruxton, Thomas & Wright, 2002). Collectively, however, cat populations
may kill very large numbers of prey. For example, Woods, McDonald & Harris (2003)
estimated that pet cats in Britain killed 52–63 million mammals, 25–29 million birds and 4–
6 million reptiles and amphibians during a 5-month survey period. However, despite the fact
that large numbers of prey are certainly taken, the impact of predation on prey populations
is equivocal (e.g. Barratt, 1997, 1998), and is dependent on the degree to which cat predation
is additive or compensatory to other sources of mortality.
With the continued expansion of urban areas into rural habitats, the importance of cat
predation as an ecological factor is potentially increasing. Urban areas currently occupy up
to 10% of the total land area of Britain and Ireland (Haines-Young et al., 2000), with around
6000 ha of rural land converted to urban use in Britain annually (Department of Environ-
ment, Food and Rural Affairs, 2003). For a number of bird species, urban populations
comprise a substantial component of the total national population (Bland, Tully & Green-
wood, 2004), including some that are of conservation concern, e.g. house sparrow Passer
domesticus, starling Sturnus vulgaris. Conversely, cat predation may be beneficial where pest
species, such as common rat Rattus norvegicus and house mouse Mus domesticus, are taken.
At present, there are few data on the pattern and extent of predation by domestic cats in
Britain (but see Howes, 2002; Woods et al., 2003), in particular for urban areas where most
cats are likely to be present. For example, approximately 70% of UK inhabitants live in
settlements of > 10 000 residents (2001 UK National Census, Office for National Statistics,
http://www.statistics.gov.uk), although the distribution of cats is unlikely to exactly mirror
the distribution of humans as patterns of pet ownership are likely to be influenced by where
people live. Therefore, we undertook a questionnaire survey of householders within north-
west Bristol, UK, to (i) quantify cat density and (ii) quantify the number of prey killed, by
asking cat owners to record prey animals returned home during each of four survey periods.
The potential impact of predation was then estimated by comparing annual predation rates
with published estimates of prey density and productivity.

METHODS
A questionnaire was delivered to all 3494 householders in a 4.2-km2 area in north-west Bristol
during October – December 2002. This is an area of predominantly semidetached housing
built during the interwar years, but also comprises regions of private flats, terraced and
detached housing, and council housing; other major habitats include sports fields, woodland,
churchyards and a cemetery (see Harris, 1981). The questionnaire requested: the number of
pet cats owned; each cat’s sex and age; whether they had been neutered; whether they wore
an antipredation device (bell, sonic emitter); whether they were typically allowed out during
the day only, during the night only or during both the day and night; and to estimate the
approximate number of birds, mice/voles and amphibians/reptiles that each cat brought home
each month. Finally, each owner was asked whether he/she would assist with the study by
keeping a simple diary recording the prey items the cat(s) brought home.
We initially asked householders to leave the completed form on their doorstep on a given
day for collection (hereafter termed ‘original’ questionnaires). Following this, a subset of
householders that failed to return the form were then contacted door-to-door and asked to

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304 P. J. Baker et al.

complete the form (‘follow-up’ questionnaires). In all analyses, original and follow-up ques-
tionnaires were first compared to determine if the two data sets were statistically different.
In those cases where differences were detected, the mean values from the follow-up question-
naires were used to extrapolate the responses of non-respondents, assuming that follow-up
questionnaire results were more representative of non-respondents.
All cat owners who stated that they would keep a diary were contacted and asked to record
the number of dead and living prey items brought in by their pet(s) during 5 December 2002
– 13 January 2003 (winter: n = 40 days), 28 April – 1 June 2003 (spring: n = 35 days), 4
October – 2 November 2003 (autumn: n = 30 days) and 24 July – 31 August 2004 (summer:
n = 39 days) inclusive. Householders were asked to report all dead animals brought in by their
cat(s) so that they could be collected for identification: prey recorded as, e.g. ‘mouse’ or ‘small
bird’, and not identified by us, were classified as unidentified. Sampling periods were spaced
out throughout the year in an attempt to reduce participant dropout. At the end of each
sampling period, non-respondents were contacted up to five times in an attempt to minimize
biases arising from, for example, the disproportionate loss of owners whose cats failed to
return any prey during the sampling period. Where possible, owners were asked to identify
which cat brought in each prey item.

Estimating predation rates

The potential impact of cat predation on prey populations was estimated by comparing the
minimum, intermediate and maximum number of animals killed annually with published
estimates of prey abundance and productivity. Predation rates were based only on those
animals returned home dead. Minimum predation rates in each season were calculated as the
total number of identified dead prey returned divided by the total number of cats in the survey
period. Intermediate and maximum predation rates were derived by first classifying uniden-
tified prey, assuming that they exactly mirrored the distribution of identified prey. It is
possible that householders were most likely to be able to identify common prey species, such
that unidentified prey would have tended to consist of other less common species: redistrib-
uting these less readily identified prey would therefore have the effect of elevating true rates
of predation on common species. Alternatively, householders may have simply been unable
to identify species generally, in particular juvenile birds: in this case, redistributing unidenti-
fied prey would reflect rates of predation on common species. Unidentified mammal and bird
remains were redistributed among identified small mammal and passerine categories (i.e.
excluding grey squirrel Sciurus carolinensis and columbiforms). Intermediate predation rates
were then calculated by dividing the number of dead prey returned by the total number of
cats. Maximum predation rates were calculated by dividing the number of dead prey returned
by the total number of cats that returned ≥ 1 prey, including instances where, for example,
only one prey was recorded for a household containing two cats. All predation rates were
then multiplied by a factor of 3.3: Kays & DeWan (2004) recorded a return rate of 1.67 prey/
cat/month (n = 12 cats) using a similar approach to that adopted in the current study, com-
pared with a kill rate of 5.54 prey/cat/month for 11 radio-collared cats whose hunting
behaviour was observed directly. The number of prey killed annually per unit area (km2) was
then calculated by multiplying the predation rate in each corresponding survey period by 91
days and summing across the year.

Potential impact of cat predation

Pre-breeding small mammal density was derived from the adjusted density estimates for
woodland (9% of study area), scrub (24%), gardens of semidetached houses (20%) and

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Predation by domestic cats 305

gardens of detached houses (8%) quoted by Dickman & Doncaster (1987): data quoted for
semidetached houses and woodlands were utilized to estimate density in the gardens of
terraced houses (5%) and allotment gardens (1%), as these habitats appear comparable
(Ansell, 2004). All other habitats were assumed to be unproductive. Grey squirrel density was
taken as 0.1/ha (Harris et al., 1995). Common rat density was estimated assuming that 13%
of households were infested (Langton, Cowan & Meyer, 2001), with a mean of 2.2 rats per
infestation (Harris et al., 1995).
Maximum productivity was estimated from the length of the breeding season divided by
the length of each breeding attempt (gestation plus lactation) multiplied by average litter size
(Corbet & Harris, 1991), i.e. ignoring any effect of density on the proportion of females
breeding. For rats, it was assumed that 30% of breeding females produced a maximum of five
litters throughout the year (Corbet & Harris, 1991): this is equivalent to each female produc-
ing 1.5 litters/year.
Breeding bird density in the City of Bristol has been measured annually in selected 1 km
squares as part of the British Trust for Ornithology’s Breeding Bird Survey for a number of
years, and during 2001, over 58% of the city was surveyed (Bland, 2001). In each survey
square, two 1-km transects were walked early in the morning twice each year, first between
1 April – 15 May, and then again at least 4 weeks later before the end of June. Surveyors were
able to recognize species by sight and song. Average densities of species in the nine 1 km
squares centred on the study site (OS grid co-ordinate: 356176) were obtained from R. Bland
and J. Tully (personal communication). Productivity was estimated by multiplying prey
density by the number of broods raised and average brood size at fledging (http://
www.bto.org). For both birds and mammals, the pre-breeding population sex ratio was
assumed to be equal.
The minimum impact of cat predation was calculated by dividing the minimum number
of animals killed by the sum of pre-breeding density and productivity. As the number of
offspring produced annually typically exceeds the size of the pre-breeding density, maximum
impact was calculated by dividing the maximum number of animals killed by pre-breeding
density alone. Intermediate impacts (ranging from smallest to largest) were calculated by
dividing the intermediate number of animals killed by pre-breeding density and productivity
combined, productivity alone and pre-breeding density alone, respectively. In all cases, we
have assumed that live individuals rescued by owners and released would have survived.

RESULTS
Overall, 1243 questionnaires (36%) were collected (1027 original, 216 follow-up). There was
a significant difference between the proportion of ‘original’ (24%) and ‘follow-up’ (17%)
respondents owning cats (χ21 = 4.64, P < 0.05). Assuming that the same proportion of non-
respondents on the study site owned cats as in the follow-up survey, it was estimated that 674
householders (19%) owned cats on the study site. There was no significant difference between
original and follow-up respondent cat owners with respect to the mean number of cats they
owned (Mann–Whitney test: W = 4458.5, N1 = 246, N2 = 37, NS; mean = 1.43 cats/house-
hold). Overall, cat density was 229 cats/km2, with a total population of 962 cats (28 cats per
100 households).
Eighty-nine householders (n = 131 cats) participated in data collection, although not all
householders collected data in all seasons. Overall, 271 dead and 87 living prey items were
recorded (Table 1). The most commonly recorded prey species was the wood mouse Apode-
mus sylvaticus, accounting for 62% and 82% of records of dead and living prey, respectively.
Estimated intermediate predation rates for each season were: spring, 1348 prey/km2; summer,

© 2005 Mammal Society, Mammal Review, 35, 302–312

 Table 1. Summary of prey items brought home

Spring Summer Autumn Winter Total

(n = 121 cats) (n = 92 cats) (n = 102 cats) (n = 117 cats) (n = 131 cats)
(n = 81 houses) (n = 63 houses) (n = 72 houses) (n = 80 houses) (n = 89 houses)
(n = 35 days) (n = 39 days) (n = 30 days) (n = 40 days) (n = 144 days)
306 P. J. Baker et al.

Species Dead Living Dead Living Dead Living Dead Living Dead Living

Mammals
Common shrew Sorex araneus 2 [2.3] 0 0 [0.0] 0 0 [0.0] 0 0 [0.0] 0 2 0
Grey squirrel Sciurus carolinensis 0 [0.0] 0 0 [0.0] 0 2 [2.0] 0 2 [2.0] 0 4 0
Wood mouse Apodemus sylvaticus 28 [32.1] 14 54 [54.0] 25 51 [51.9] 17 35 [36.9] 15 168 71
Bank vole Clethrionomys glareolus 1 [1.1] 0 5 [5.0] 0 0 [0.0] 0 0 [0.0] 0 6 0
Common rat Rattus norvegicus 3 [3.4] 1 1 [1.0] 0 1 [1.0] 1 1 [1.1] 1 6 3
Unidentified rodent 3– 1 0– 0 0– 0 1– 0 4 1
Unidentified mammal 2– 0 0– 0 1– 0 1– 0 4 0
Birds
House sparrow Passer domesticus 8 [11.4] 3 6 [9.0] 0 0 [0.0] 0 0 [0.0] 1 13 4
Dunnock Prunella modularis 4 [5.7] 0 1 [1.5] 0 1 [2.0] 0 0 [0.0] 0 2 0
Blue tit Parus caeruleus 3 [4.3] 0 2 [3.0] 0 0 [0.0] 0 3 [3.4] 0 8 0
Great tit Parus major 1 [1.4] 0 0 [0.0] 0 0 [0.0] 0 0 [0.0] 0 1 0
Blackbird Turdus merula 2 [2.8] 0 0 [0.0] 0 0 [0.0] 0 2 [2.3] 0 4 0
Starling Sturnus vulgaris 3 [4.3] 0 0 [0.0] 0 0 [0.0] 0 0 [0.0] 0 1 0
Robin Erithacus rubecula 8 [11.4] 0 3 [4.5] 0 0 [0.0] 0 1 [1.1] 0 7 0
Wren Troglodytes troglodytes 2 [2.8] 0 0 [0.0] 0 0 [0.0] 0 1 [1.1] 0 3 0
Wood pigeon Columba palumbus 0 [0.0] 0 0 [0.0] 0 0 [0.0] 0 1 [1.0] 0 1 0
Feral pigeon Columba livia 0 [0.0] 0 1 [1.0] 0 2 [2.0] 0 0 [0.0] 0 3 0
Unidentified bird 13 – 4 6– 2 1– 0 1– 0 33 6
Herpetofauna
Common frog Rana temporaria 1 [1.0] 2 0 [0.0] 0 0 [0.0] 0 0 [0.0] 0 1 2
Total 84 [84.0] 25 79 [79.0] 27 59 [59.0] 18 49 [49.0] 17 271 87

Data in square brackets indicate revised seasonal totals after redistribution of unidentified prey remains (see text for details).

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Predation by domestic cats 307

1514 prey/km2; autumn, 1324 prey/km2; and winter, 720 prey/km2 (Table 2). This was equiv-
alent to an annual rate of 4906 prey/km2 or 21 prey/cat/annum. However, in all seasons, the
majority of cats (57%, 51%, 73% and 74%, respectively) did not return any dead prey
(Table 2). Pre-breeding prey abundance and productivity is summarized in Table 3, and the
potential impact on prey populations is outlined in Table 4.
Fifty-two households, containing 77 cats, recorded data in all four seasons. Collectively,
these returned 212 dead prey items, an average of 2.8 prey items per cat in the sampling
periods (n = 144 days) or 7.0 prey/cat/annum. There was no significant difference between
seasons in the predation rate per household for this subsample (Friedman test: χ23 = 5.64,
P > 0.05), although a large number of these households had zero rates for three (n = 9
households) or four (n = 17) seasons.

DISCUSSION
The intermediate predation rate observed in this study (21 prey/cat/annum) was at the low
end of the spectrum reported in similar previous studies in the UK (14 – Churcher & Lawton,
1987; 28.9 – Howes, 2002; 71.7 – Ruxton et al. (2002), based on an average of 5.5 prey per
cat in a sampling period of 28 days; and 27.0 – Woods et al. (2003), based on an average of
11.3 prey per cat in a sampling period of 153 days), principally because in each sampling
period, the majority of cats (51–74%) failed to return any prey. However, there are a number
of problems inherent with drawing direct comparisons between studies.
First, the predation rates determined in this study were most reliant on cat density, as this
parameter had the greatest magnitude (229 cats/km2). We adopted a two-sampling protocol
to adjust for the fact that cat owners may have been more likely to leave the original
questionnaire on the doorstep as requested. Yet, the follow-up survey could still have over-
estimated cat density if some householders were more likely to have been absent when we
called and these absent householders were less likely to own cats. However, given that the
major demographic groupings on the study site were pensioners and middle-class families
with children of school age, and given that we attempted to contact residents both during
the day and in the evening, it is our opinion that the magnitude of any bias against non-cat-
owning households is likely to be minimal.
Second, recruiting participants into surveys is also susceptible to biases. For example,
owners whose cats kill large numbers of prey may be over-represented if owners are keen to
demonstrate their pet’s hunting prowess and/or if owners of cats that kill very few or no prey
consider that such information is not important. We attempted to control for this problem
by including a section on the original questionnaire asking householders to estimate the
approximate number of prey that each of their cats brought home each month. These data
could then have been used to estimate whether the composition of participating owners was
biased towards cats that tended to bring large numbers of prey home (which would have
overestimated the true pattern of predation) or that tended to bring no prey home (which
would have underestimated predation rates). Unfortunately, the vast majority of cat owners
did not answer this question or gave information that could not be readily quantified.
Consequently, we considered this comparison unreliable. Yet, it must be noted that the
majority of cats in each survey period did not bring home any prey, and that half of the
households providing year-round data had zero counts in three or four seasons, indicating
that the estimated predation rates are not likely to be excessive in magnitude. This is partic-
ularly true if cats that did not return any prey were actually taking prey (see below). Predation
rates would also be higher if, as seems probable, some live prey individuals rescued by owners
subsequently perished.

© 2005 Mammal Society, Mammal Review, 35, 302–312

 Table 2. Estimated minimum (min.), intermediate (int.) and maximum (max.) seasonal predation rates (prey killed per km2)

Spring Summer Autumn Winter

Annual intermediate
Min. Int. Max. Min. Int. Max. Min. Int. Max. Min. Int. Max. predation rate
308 P. J. Baker et al.

Species (n = 121) (n = 121) (n = 52) (n = 92) (n = 92) (n = 45) (n = 102) (n = 102) (n = 28) (n = 117) (n = 117) (n = 30) (minimum–maximum)

Mammals
Common shrew 32 37 87 0 0 0 0 0 0 0 0 0 37 (32–87)
Grey squirrel 0 0 0 0 0 0 45 45 164 29 31 120 76 (74–284)
Wood mouse 455 521 1213 1035 1035 2116 1146 1166 4249 514 542 2115 3265 (3150−9692)
Bank vole 16 18 42 96 96 196 0 0 0 0 0 0 114 (112–237)
Common rat 49 55 128 19 19 39 22 22 82 15 16 63 113 (105–313)
Birds
House sparrow 130 185 431 115 172 353 0 0 0 0 0 0 358 (245–783)
Dunnock 65 93 215 19 29 59 22 45 164 0 0 0 166 (107–438)
Blue tit 49 70 162 38 57 118 0 0 0 44 50 195 177 (131–475)
Great tit 16 23 53 0 0 0 0 0 0 0 0 0 23 (16–53)
Blackbird 32 45 106 0 0 0 0 0 0 29 34 132 79 (62–238)
Starling 49 70 162 0 0 0 0 0 0 0 0 0 70 (49–162)
Robin 130 185 431 57 86 176 0 0 0 15 16 63 288 (202–670)
Wren 32 45 106 0 0 0 0 0 0 15 16 63 62 (47–169)
Wood pigeon 0 0 0 0 0 0 0 0 0 15 15 57 15 (15–57)
Feral pigeon 0 0 0 19 19 39 45 45 164 0 0 0 64 (64–203)
Total 1055 1348 3136 1399 1514 3096 1281 1324 4822 676 720 2808 4906 (4412−13862)

Minimum predation rates were determined by dividing the number of identified prey by the total number of cats in each season. Maximum predation rates were determined
by dividing the total number of dead prey (unidentified prey redistributed in accordance with distribution of identified prey) by the number of cats returning ≥1 prey in each
season. Intermediate rates were determined by dividing the total number of dead prey by the total number of cats in each survey period. Figures in brackets indicate the number
of cats used in each calculation. Rates were then multiplied by a factor of 3.3 to account for the proportion of prey killed that was returned home (see text for details). Each
season was assumed to last 91 days. Note that subtotals for minimum estimates excluded unidentified prey and are therefore not true totals.

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Predation by domestic cats 309

Table 3. Estimated patterns of prey density and productivity (see text for details)

Pre-breeding Productivity (young per year)

density
Species (prey/km2) Litter/brood per year Litter/brood size Annual production

Mammals
Common shrew 1382 3.8 6.0 15 755
Grey squirrel 100 1.6 3.0 240
Wood mouse 2051 5.1 5.5 28 770
Bank vole 623 4.6 3.6 5 162
Common rat 238 1.5 7.5 1 338
Birds
House sparrow 120 2.5 2.84 426
Dunnock 50 2.0 3.67 184
Blue tit 120 1.0 7.50 450
Great tit 60 1.0 6.56 197
Blackbird 180 2.5 3.36 756
Starling 60 1.5 3.86 174
Robin 80 2.0 4.50 360
Wren 100 1.0 5.16 258

Table 4. Comparison of the minimum (m), intermediate (I) and maximum (M) predation rates (Table 2) with
pre-breeding prey abundance (A) and productivity (P) (Table 3)

Impact

Intermediate
Predation rate Minimum Maximum
Species (prey/km2/year) {m/(A + P)} {I/(A + P)} {I/P} {I/A} {M/A}

Mammals
Common shrew 37 (32–87) <0.01 <0.01 <0.01 0.03 0.06
Grey squirrel 76 (74–284) 0.22 0.22 0.32 0.76 2.84
Wood mouse 3265 (3150−9692) 0.10 0.11 0.11 1.59 4.73
Bank vole 114 (112–237) 0.02 0.02 0.02 0.18 0.38
Common rat 113 (105–313) 0.07 0.07 0.08 0.48 1.31
Birds
House sparrow 358 (245–783) 0.45 0.65 0.84 2.98 6.53
Dunnock 166 (107–438) 0.46 0.71 0.91 3.33 8.76
Blue tit 177 (131–475) 0.23 0.31 0.39 1.48 3.96
Great tit 23 (16–53) 0.06 0.09 0.12 0.38 0.88
Blackbird 79 (62–238) 0.07 0.08 0.10 0.44 1.32
Starling 70 (49–162) 0.21 0.30 0.40 1.16 2.71
Robin 288 (202–670) 0.46 0.65 0.80 3.59 8.38
Wren 62 (47–169) 0.13 0.17 0.24 0.62 1.69

Third, studies of cat predation are drawn from different habitats with different prey bases,
such that predation rates of cats in rural areas with increased levels of available prey may be
substantially higher (e.g. Howes, 2002). The relatively small number of prey species taken
during this study is consistent with the reduction in species’ diversity typically observed in
urban areas, where communities tend to consist of a small number of synanthropic species
tolerant of human activities, but which may be very abundant (Luniak, 2004). In high-density
predator populations, such as in this study, individuals may also be restricted to small home
ranges (Kays & DeWan, 2004), thereby reducing the range of prey they can encounter and
their impact, but which may enhance localized effects (see Baker et al., 2003).

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310 P. J. Baker et al.

Last, the method utilized implicitly assumes that the rate at which prey is returned home
is consistent across populations, yet the magnitude of any difference between populations is
unknown. In this study, we have utilized a conversion factor of 3.3 to estimate the actual
number of prey killed, based on the work of Kays & DeWan (2004). To the best of our
knowledge, this is the only study that has managed to quantify this parameter. Consequently,
there is the need for additional studies to determine the consistency of this estimate. Further-
more, attention must also be paid to cats that fail to return any prey home; the assumption
that these individuals had not killed anything would have significantly underestimated pre-
dation rates if they were taking prey.

Impact of cat predation

Minimum predation rates were moderately high for three species: house sparrow (predation
equivalent to 45% of the combined total of pre-breeding density and annual productivity),
dunnock (46%) and robin (46%: Table 4). In these species, intermediate predation rates were
equivalent to 80–91% of productivity alone and 65–71% of the combined total of pre-
breeding density and annual productivity. Such losses are likely to be non-trivial and warrant
further investigation, in particular whether cat predation is additive or compensatory to other
forms of mortality.
The difference in magnitude of these estimates is dependent on the treatment of unidenti-
fied prey remains. Minimum rates were derived solely from identified prey items, whereas
intermediate rates were estimated from both identified and unidentified prey remains under
the assumption that unidentified prey consisted of the same species, and in the same relative
abundance, as those that had been identified. We believe that this was likely to be true since
the majority of unidentified prey probably related to predation on juvenile birds in spring
and summer, which householders were unlikely to be able to identify even if they were
common species. The assumption of identical distributions is less clear, and future studies
should therefore aim to recover all carcasses for positive identification.
The house sparrow has previously been identified as potentially vulnerable to the impact
of cat predation (Churcher & Lawton, 1987), and is typically the most common avian species
taken in UK studies (Howes, 2002; Woods et al., 2003; but see Mead, 1982). Nationally, this
species is declining in urban areas (Siriwardena, Robinson & Crick, 2002), but numbers
appear to be stable in Bristol as a whole (J. Tully, personal communication). However, given
that house sparrow distribution is highly heterogeneous and that the study site was an area
of low density (Bland, 1998), it is possible that cat predation was significantly affecting levels
of recruitment and creating a dispersal sink for more productive neighbouring areas: similar
effects may also have been evident for robins and dunnocks. Consequently, we would recom-
mend quantification of age-specific patterns and rates of mortality, predation and dispersal
in these species.
Overall, therefore, the levels of predation by individual cats observed in this study were
towards the lower end of rates quantified in previous studies, yet direct comparisons between
studies are difficult. Collectively, despite occurring at very high densities, the summed effects
on prey populations appeared unlikely to affect population size for the majority of prey
species. This may be attributable to selection for prey species that have successfully adapted
to the urban environment, including the predation pressure exerted by cats. However, for
three species there was evidence that the impact of cat predation may be significant. Conse-
quently, the overall impact of predation by domestic cats in urban areas warrants further
investigation, including: (i) consideration of prior species’ loss in the formation of current
urban animal communities; (ii) the use of manipulative experiments and/or comparative

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Predation by domestic cats 311

studies to quantify the effects of reduced predation pressure on prey population growth; (iii)
quantifying the relative importance of cat predation within the guild of mammalian and avian
predators present in urban areas; and (iv) monitoring of the relative importance of cat
predation in different urban habitats and over time.

ACKNOWLEDGEMENTS
We would like to express our gratitude to: Philippa Bulman, Eve Jackson, Louise Jasper,
Fiona Strouts, Stephanie Thein and members of the general public for help in collecting the
data; Richard Bland and John Tully for providing data on bird densities; and Stuart Newson
and colleagues at the British Trust for Ornithology for help in identifying bird carcasses.

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Submitted 8 February 2005; returned for revision 11 May 2005; revision accepted 1 July 2005
Editor: RM

© 2005 Mammal Society, Mammal Review, 35, 302–312