Full Terms & Conditions of access and use can be found athttp://www.tandfonline.com/action/journalInformation?journalCode=tnzz20

Download by: [119.224.56.228] Date: 06 November 2016, At: 23:20

New Zealand Journal of Zoology

ISSN: 0301-4223 (Print) 1175-8821 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzz20

Seventeen years of predation by one suburban catin New Zealand

John E. C. Flux

To cite this article: John E. C. Flux (2007) Seventeen years of predation by onesuburban cat in New Zealand, New Zealand Journal of Zoology, 34:4, 289-296, DOI:10.1080/03014220709510087

To link to this article: http://dx.doi.org/10.1080/03014220709510087

Published online: 19 Feb 2010.

Submit your article to this journal

Article views: 1305

View related articles

Citing articles: 9 View citing articles

New Zealand Journal of Zoology, 2007, Vol. 34: 289-2960301-4223/07/3404-0289 © The Royal Society of New Zealand 2007

Seventeen years of predation by one suburban cat in New Zealand

JOHN E. C. FLUXEcological Research Associates of New Zealand230 Hill RoadBelmontLower Hutt, New Zealandflux@paradise.net.nz

Abstract The 558 prey items brought home byone domestic cat, recorded over its 17-year lifetime,included 221 mice, 63 rats, 35 rabbits, 4 hares and2 weasels. The cat hunted up to 600 m from thehouse, and prey was caught both inside and outsidethe 0.5 ha garden. Of the 223 birds brought in, 54were native, including 43 silvereyes (Zosterops lat-eralis), but those killed were quickly replaced, sothere was always a resident population of 1-2 pairs.The other known native birds comprised five fantails(Rhipidura fuliginosa), four warblers (Gerygoneigata), a kingfisher (Halcyon sancta), and a shiningcuckoo (Chrysococcyx lucidus). Only nine skinks(Cyclodina aenea) and one frog (Litoria raniformis)were brought in. The abundance of birds and reptilesin the garden showed no apparent change over the17 years compared with the previous 15-year-periodwithout a cat. By contrast, the cat exterminated therabbit population in the garden, and "farmed" sur-rounding burrows during its whole life; all otherprey killed declined in frequency after the cat was8-9 years old.

Keywords birds; domestic cat; Felis catus; mus-telids; predation; rabbits; rodents

INTRODUCTION

Since the pioneering studies of the prey broughthome by domestic cats (George 1974; Borkenhagen1978; Churcher & Lawton 1987), there has been a

Z07015; Online publication date 14 September 2007Received 22 April 2007; accepted 25 August 2007

growing appreciation of the possible importance ofdomestic cats as predators of urban wildlife in sev-eral countries, including Australia and New Zealand(e.g., Barratt 1997, 1998; Finnegan 1992; Gillies &Clout 2003).

Most studies of the diet of domestic cats havebeen short-term (1-2 years) and either recorded theprey of individual cats or averaged the kills of a largenumber of cats from data submitted by their owners(see review by Fitzgerald & Turner 2000). Analysesof kills by cats of different ages indicate a decline inthe number of kills with age (Barratt 1998; Churcher& Lawton 1987; and many subsequent questionnairestudies), but Fitzgerald & Turner (2000) note that forindividual cats, changes in diet with age are poorlydocumented. Hence, this first report documenting thevariation in prey numbers taken by an individual catover an entire life span is of interest, although notnecessarily representative of cats in general.

METHODS

Study areaThe study area was a 0.5 ha garden in the hills aboveLower Hutt (41°11'S, 174°55'E). It has a mature 10ha Pinus radiata plantation on one side, and opengrassland with scattered gorse (Ulex europaeus)and scrub on the other. In 1972, the plantation was20 years old with a closed canopy and little under-growth; since then it has become far more open,with regenerating native species (mahoe Melicytusramiflorus, fivefinger Pseudopanax arboreus, andwineberry Aristotelia serrata) reaching 10 m inheight. The grassland had been closely grazed bysheep, but was then leased to a pony club and grazedby horses. Although cattle and sheep were broughtin to control weeds, the grass became rougher andpatches of gorse grew.

The garden has also changed from a relativelywell-tended mix of lawn, roses, hydrangeas andother introduced garden plants to a rougher, morenatural style with tree ferns, wineberry, mahoe,

2W

290 New Zealand Journal of Zoology, 2007, Vol. 34

Fig. 1 The cat was able to climbon thin twigs, but mostly huntedon the ground.

fivefinger and pepper tree Macropiper excelsumregenerating over about a quarter of the area. Halfthe garden is in rough grass with fruit trees.

FaunaMice (Mus musculus) and rats (only Rattus rattus)lived in sheds and around compost bins, and both en-tered the house every autumn. Rabbits were residentand breeding in the garden from 1972-88, but abouttwice as many hares (Lepus europaeus) as rabbits(Oryctolagus cuniculus) lived in the field oppositethe house (Flux in press). Possums (Trichosurusvulpecula) were always present in low numbers de-spite sporadic trapping to protect fruit or ornamentaltrees (removing an average of 21 (0-54) possumsa year over 27 years). Stoats (Mustela erminea)were seen every 2 or 3 years; one ferret (Mustelaputorius) was trapped. Intensive mustelid trappingin the adjacent Belmont Regional Park (Flux & Bull1976) yielded no weasels (Mustela nivalis), but thecat caught two.

Birds in the garden were observed regularly from1972, but not fed. However, house sparrows (Passerdomesticus), hedgesparrows (Prunella modularis)and blackbirds (Turdus merula) ate the hens' spiltmash, and a trap baited with fat for catching sil-vereyes for banding, set in winter from 1972 to 1976,also caught house sparrows and finches. Any unusualspecies was recorded, and the numbers of residentbirds estimated from observations and nests foundeach year. The resident breeding populations of thecommon small birds included house sparrows, 5-10pairs (except in 2000-01 when local sparrows were

eliminated by disease); hedgesparrows, 2-3 pairs;starlings (Sturnus vulgaris), 2-7 pairs (Flux 2003);chaffinch Fringilla coelebs, greenfinch Carduelischloris, goldfinch C. carduelis and yellowhammerEmberiza citrinella, 1-2 pairs each; blackbirds 3-5pairs; song thrushes 1-2 pairs. Native birds includ-ing silvereyes, fantails and warblers were relativelytame and present throughout the year. Copper skinkswere common in the garden but were most often seenwhen caught by the resident kingfisher. The gardenwas also free range to 6-15 hens and a rooster.

The cat (Fig. 1) was a neutered female born inthe wild towards the end of 1987. She was fed adlib. morning and night, but never became fat, weigh-ing 3.5 kg for most of her life. In her last year shedeveloped a lung problem, and when put down on2 May 2005 weighed 2.8 kg.

Prey recordingAll the specimens were collected and identified bya zoologist, thus avoiding the uncertain identifica-tion common in public questionnaires. Many werebrought into the house, or left on the doorstep, butothers were seen as the cat ate them in the garden orcarried them past the house. A few specimens thatwere not found fresh were allocated a date based onappearance and smell. As some kills would not havebeen found, and we were overseas occasionally, thefigures are minimum estimates; but there was noreason to expect bias in the ratio of species found.Three empty bird nests brought to the door had ap-parently held chicks that had been eaten: goldfinch,3 January 1990), chaffinch, 20 December 1991)

Flux—Seventeen years of predationby suburban cat 291

and greenfinch, 22 December 1998). These are notincluded in later analyses, but seem to be the firstrecord of such behaviour.

Although the cat often killed and ate insects, deadinsects could not be identified as cat kills and are notincluded in this analysis.

RESULTS

Hunting methodsOnly two successful hunts of vertebrates were seen.One was a leap from a 1.2 m fence post, landing 2 maway on a mouse in grass 30 cm high; and a youngrabbit was jumped on as it ran in a ditch. For boththe cat relied entirely on hearing. She often playedwith mice and fledgling birds before killing them.One skink escaped while the cat pawed its cast tail.A stoat carrying a mouse was chased under awood-pile, and the cat emerged with the mouse.

She climbed trees up to 15 m high readily (Fig. 1),and often inspected two starling nest-boxes in thegarden. She entered hollow, standing, tree-ferntrunks head first, and reversed out.

Prey takenThe list of prey identified (Table 1), shows mam-mals were the main diet (58.2%) followed by birds(40.0%) and herps (1.8%). Of the larger mammals,more juveniles were brought home: e.g., four haresabout a week old (100-150 g), and, not surprisingly(since a rabbit will produce four times as many youngin a year as a hare, and young hares are notoriouslydifficult to find) 34 young rabbits of 200-250 g,plus one adult. Some rabbit burrows were occupiedfor over 20 years, others dug afresh near those ofprevious years. The cat checked them regularly andknew when the young emerged, waiting for hoursat the burrow entrance. She seemed to "farm" therabbits, and brought in young at monthly intervalsas the female produced them (Flux 1997).

Table 1 Age of prey taken by cat.

Mammals

MouseWeaselRatRabbitHareTotalBirds

HedgesparrowSilvereyeGreenfinchGoldfinchChaffinchHouse sparrowBlackbirdThrushYellowhammerStarlingFantailWarblerKingfisherShining cuckooRedpollUnidentified chicks

TotalHerpsSkinkFrogTotal

Totals

Adult

2022

2310

228Adult

294014•9

1314

9

3

.34111

144

91

10(1.8%)

382

Juvenile

19

40344

97Fledgling

17

an1232443

.2

61

158

Total

2212

63354

325 (58.2%)Nestling

2

511

16

•2

18

18

Total

48433022171613776541112

223 (40.0%)

558

292 New Zealand Journal of Zoology, 2007, Vol. 34

Most of the resident species were recorded as catkills, except any large birds like kereru Hemiphaganovaeseelandiae, mallardAnas platyrhynchos, para-dise shelduck Tadorna variegata, Australasian harrierCircus approximates, Australian magpie Gymnorhinatibicen, white-faced heron Ardea novaehollandiae,spur-winged plover Vanellus miles, sulphur-crestedcockatoo Cacatua galerita, eastern rosella Platycer-cus eximius, morepork Ninox novaeseelandiae andblack-backed gull Larus dominicanus that landedin the cat's hunting area. The relative proportions ofsome species brought in, such as house sparrows andhedgesparrows (16:48, Table 1) did not match theirapparent availability; others, such as blackbirds andsong thrushes, appeared as kills in about the sameproportion (3:1) as they were observed alive. Thecat never molested the hens or their chicks.

Overall, 137 adult birds were killed comparedwith 77 nestlings plus fledglings, and even dur-ing the months in which young birds were taken(September-March) the totals still favoured adults91:11. Only from November to January were adultsoutnumbered (40:56).

Adult silvereyes, house sparrows and chaffincheswere far commoner prey than fledglings of thesespecies, and only one nestling was killed among75 adults. Other species like hedgesparrow, green-finch, goldfinch, blackbird, song thrush (Turdusphilomelos) andyellowhammer showed a more evenbalance between adult and fledgling kills. Consid-erably more fledglings than nestlings were caught(61 cf. 16), even though nestlings were availablefor 10-15 days in the nest, compared with less thanhalf that time as a fledgling (defined for this studyas a young bird capable of flight but still unlikely to

have left the nest). Of 54 native birds recorded, 43were silvereyes. With the exception of silvereyes,native birds such as fantails and warblers, whichare relatively tame and present throughout the year,were seldom recorded.

Copper skinks brought in were played with likemice, making them easy to record. No geckos (Hop-lodactylus maculatus) were killed. (A new cat caughtone in June 2006.) A full-grown green frog (none hadever been seen or heard in the garden, which has noponds) was carried into the house, but then dropped,while the cat foamed at the mouth.

Intensity of hunting with ageThe number of prey items brought in by the catincreased during the first 4 years of her life, andreached a plateau of over 60 items per year at age3-7 (Table 2). From 8 to 12 years old there was adecline to 15 items; and in her final 5 years, aged13-17, she averaged only five items per year. Thepattern of increase, plateau and decline seems to bemirrored in all the prey species except rabbits, whichwere taken whenever available, until the cat was 16.There was no swing from birds to mammals, whichmight be easier for an older cat to catch; indeedthe ratio of introduced birds to mammals increasedslightly from 1:2.03 at ages 1-9 years, to 1:1.77 atages 10-17 years. The proportion of native to intro-duced birds declined significantly (Chi square 8.6, P= 0.01) from 1:2.6 to 1:30 over the same periods.

Seasonal distribution of killsThere was a pattern of increased activity in summerand autumn for most species brought in by the cat(Table 3).

Table 2 Number of kills made by the cat during its lifetime.

Age of cat

MiceRatsRabbitsHaresWeaselsTotal

Introduced birdsNative birdsTotal

SkinksFrogTotal

Totals

1

03

Is)

5

4

Is)

6

11

Is)

16

Is)

Is)

1

21

11

Is)

15

34

3

2010

Is)

1

33

267

33

1

1

67

4

3393

45

181230

Is)

13

78

5

29103

143

179

26

Is)

Is)

71

6

374

Is)

144

236

29

Is)

Is)

75

7

355

40

171025

67

8

851

14

163

19

33

9

1638

27

7

Is)

9

Is)

Is)

38

10

1157

23

5

5

28

11

83

11

5

5

16

12

Is)

Is)

Is)

6

819

15

13

11

Is)

4

1

1

5

14

Is)

1

3

7

7

10

15

1

Is)

3

1

1

4

16

1

1

Is)

Is)

Is)

4

17

1

1

1

1

2

Flux—Seventeen years of predationby suburban cat 293

Proportion of each carcass eatenFrom 1992 the condition of the prey was recorded:91 mice were brought in live or found dead uneaten;31 were partly eaten, and 23 completely eaten. Ofthe rats, only 10 were uneaten, 15 partly eaten, and8 completely eaten. Rabbits, however, were alwayseaten entirely, apart from the skin and hind feet.Among the birds, 15 of 20 hedgesparrows were al-most completely eaten, compared with only 5 of 15house sparrows (Chi square = 6.1, P = 0.02). Of theother small birds, about half were eaten, including23 of 42 silvereyes and 27 of 56 finches.

DISCUSSION

Although based on the diet of only a single cat (butone born in the wild and fed as a kitten on wild-caught prey), the results reported here confirm boththe pattern found in Wellington in a much larger ifless detailed study (Gillies & Cutler 2001), and theoverall conclusion that cats are primarily predators

of small mammals, especially rabbits (Fitzgerald &Turner 2000).

Several studies have indicated that older catscatch fewer animals (see references in Fitzgerald &Turner 2000). Our cat showed a fairly rapid declinein hunting of mammals and birds from age 7. Anexception was rabbits: 15 were taken at ages 1-8,compared to 20 from age 9-16. Possibly catchingbaby rabbits requires patience rather than agility.All were eaten completely, confirming the generalopinion that rabbits are a cat's most preferred food(Borkenhagen 1978; Liberg 1984; Gibb & Fitzgerald1998). Only one of the four small hares, however,was eaten. Hunger is not a necessary condition forkilling in domestic cats (Biben 1979).

The higher numbers of most prey recorded insummer and autumn (Table 3) was expected fromtheir seasonal recruitment of inexperienced young,and greater vulnerability while nesting. The catselected fully grown fledglings rather than nest-ling chicks. Borkenhagen (1978) similarly foundthat only seven of 68 birds brought in by cats in

Table 3 Seasonal distribution of kills, 1988-2005.

MouseRatRabbitHareWeaselTotal

HedgesparrowSilvereyeGreenfinchGoldfinchChaffinchHouse sparrowBlackbirdThrushYellowhammerStarlingFantailWarblerKingfisherShining cuckooRedpollUnidentified chicksTotal

SkinkFrog

Totals

Jan

1595

Is)

132

1416

10562

2

46

4

82

Feb

23105

139

7765

Is)

1

1

1.

30

11

71

Mar

4718

65

390

Is)

Is)

4

1

1

22

87

Apr

2710

37

1501

Is)

11

11

1

49

May

233

26

005

Is)

1

Is)

Is)

1

13

39

Jun

170

17

341

1

1

10

27

Jul

164

20

164

1

12

32

Aug

93

Is)

14

100

1

15

Sep

12111

15

111

11

Is)

Is)

9

1

25

Oct

11

Is)

71

21

130

1

1

Is)

(SI

1

11

1

33

Nov

10

Is)

9

21

1150

1

43

4

28

49

Dec

1116

18

18

Is)

7Is

)

6

Is)

11

Is)

Is)

30

1

49

294 New Zealand Journal of Zoology, 2007, Vol. 34

Germany were flightless young. Mudge (2002) re-corded that two-thirds of the fantail chicks taken byowls were almost full-grown: "I have the impressionthat moreporks harvest chicks once they reach adesirable size".

Few starlings were taken despite their abundance,as Barratt (1997) also found in suburban Canberra.Cats accounted for 32% of 53 recoveries of star-lings marked in the Belmont area, mostly recentlyfledged young (Flux & Flux 1981), but our cat caughtnone.

Changes in prey populationsThe house was occupied from 1972 for 15 yearswithout a resident cat, and only a few stray catswere seen. This was followed by 17 years with theresident cat described here.

From daily observations of birds in the garden,and of nests found, an assessment of major changesin abundance could be made over these periods.Overwintering flocks of silvereyes declined from10 to 20 in 1972-76 to family parties of 3-7 birdsby 2000-05. This decline began before the cat ar-rived. The house sparrow population was eliminatedby an outbreak of salmonella in mid 2000 (the lastsparrows killed by the cat were six in early 1999)and no sparrows were seen in the garden until apair appeared on 3 September 2001. None nestedthat summer, but six were present on 12 June 2002,and they increased to the usual 5-10 pairs by thefollowing year. No other declines in resident birdpopulations were recorded, even for hedgesparrows,the birds most often brought in by the cat. Somespecies moved into the area: welcome swallows(Hirundo tahitica) in 1982, spur-winged ploversin 1987, tui (Prosthemadera novaeseelandiae) andkereru in 1991, all nesting within the hunting rangeof the cat. Tui and kereru may have been attractedby the increase in native vegetation.

Of the mammal populations present, only rabbitsshowed dramatic changes. Five pairs occupyingseparate burrows in the garden in 1987 were elimi-nated by the cat within a year, and no rabbits wereresident in the garden until 4 months after the catdied in May 2005.

Effect on native faunaThe effect of domestic cats on native fauna hasbeen much discussed in New Zealand recently, as inAmerica, Britain and Australia, (e.g., Proulx 1988;Fitzgerald 1990). Baker et al. (2005) suggest thatthree of the commonest British species may be at

risk, extrapolating from 13 (14?) house sparrows,2 (6?) hedgesparrows and 8 blue tits killed by 131cats in 144 days over 4.2 km2 in Bristol. By con-trast, Mudge (2002) found that "fantail breedingsuccess is significantly greater near houses wherethere are cats—especially when rat numbers arehigh", a statement based on a 5-year study of over400 fantail nests monitored by automatic cameras.This is not a new idea. In the 1920s, Guthrie-Smith(1953) pointed out that: "I have found by far thelarger number of Rails' nests containing unspoiledeggs close to cottages. Injurious as the cat may be tothe Rail, its presence is still more baneful to the rat;relatively that is, the cat has actually come to exerta protective influence".

The cat at Belmont took only 54 native birds, ofwhich 43 were silvereyes, a species of no conserva-tion concern. The 63 rats outnumber all the nativebirds killed, and each rat would have been capable ofkilling many birds, destroying nests and competingwith birds for food. Mice also compete with birdsfor food and may destroy a few nests, but their maindamage is likely to be to reptile populations (Ruscoe& Murphy 2005). Brockie (2001), who tallied theprey brought in by six pet cats in Wellington city,recorded 15 rats, 45 mice, 25 house sparrows, 15silvereyes, 8 blackbirds, 3 goldfinches, 3 starlings, 2fantails, 2 warblers, a hedgesparrow, a thrush, and 45lizards. He concluded that, by killing more rats andmice than native birds, the cats probably benefitedthe native bird population.

The nearest comparable study concerned 130 catsin central Wellington observed by Gillies & Cutler(2001). Most of the prey they brought in from Janu-ary to April 2000 (effectively 30 cat-years) werefound in similar proportion to our cat's kills over 17years (inbrackets): 314 (221) mice, 67 (63) rats, 267(223) birds and 1 (2) mustelids. However, two itemsdiffered: 191 (9) reptiles and 3 (35) rabbits. The rab-bits are easily explained by our rural location. Theabundant reptiles in Wellington might be a result offewer kingfishers or blackbirds, both predators oflizards (Higgins et al. 2006), as the tallies for rodentsand mustelids were similar: on the other hand, ourcat may simply have disliked them. Overall theeffect of our cat on reptiles was insignificant—thekingfisher it killed would have caught more skinksin a week than the cat's 17-year total.

Amajor difference from most other studies of citycats was the low proportion of house sparrows (8%)compared with finches (35%) and hedgesparrows(23%). Indeed, no house sparrows were caught until

Flux—Seventeen years of predation by suburban cat 295

our cat was 5 years old, despite this being the com-monest bird present. Our new cat also caught fourhedgesparrows to one house sparrow in her first year.In Britain, Mead (1982) analysed 26 889 bandingrecoveries of garden birds, and showed that 31.3%of hedgesparrows were killed by cats, the highestmortality for any species, compared with only 20.6%of house sparrows. He found no evidence of catsthreatening any species and concluded: "small birdpopulations may well be under less pressure fromcats than they would have been from the mixed bagof natural predators".

Mead's observations illustrate a problem with theuse of questionnaire methods: the public may notdifferentiate hedgesparrows from house sparrows. InWellington, Gillies & Cutler (2001) recorded 61%house sparrows and no finches or hedgesparrows(both of which are common there). InAuckland, Gil-lies & Clout's (2003) correspondents reported thathouse sparrows were the commonest bird recordedin both the urban (37%) and suburban (28%) areas;finches were "less commonly caught" and therewere no hedgesparrows. In Britain, questionnairestudies list 130 house sparrows to 14 hedgespar-rows (Churcher 1983) and 18 house sparrows to sixhedgesparrows (Baker et al. 2005), but a zoologistrecorded three house sparrows to 10 hedgesparrows(Carss 1995).

To balance the effect of various predators actingtogether is difficult. Over 50 years ago Bull (1953)warned: "It must not be forgotten that cats are onlyone of the several introduced predaceous mammalsand that these have effects on each other as wellas on the birds...even if [exterminating cats] werepossible, it would be most unwise to do so until therelationships between cats, rats and birds are morefully understood". Complex computer modellingoffers the same advice: "eradication of introducedsuperpredators, such as feral domestic cats, is notalways the best solution to protect endemic preywhen introduced mesopredators, such as rats, arealso present" (Courchamp et al. 1999). Efford et al.(2006), from a long-term study of feral cats and ratsin the Orongorongo Valley, "question the assumptionthat ship rats are 'eruptive', except possibly whenpredation is eliminated."

Advice to keep cats indoors at night (Forest &Bird 2002) is totally counterproductive in New Zea-land. They catch rodents at night, and there are nonative terrestrial mammals to protect. In Australia,Barratt (1997) found cats killed more mammals(introduced rodents) at night, and birds and reptiles

by day: keeping cats in at night was recommendedonly for new residential developments adjacent toundisturbed habitat where native mammals were atrisk (Barratt 1998). In Britain, Woods et al. (2003)found that cats kept indoors at night caught signifi-cantly fewer mammals, but there was no differencein the number of birds caught.

Outside towns, blanket control of predators iscostly to maintain in perpetuity and may also becounter-productive. For example, yellowheads (Mo-houa ochrocephala) onMt Stokes survived predationfor 150 years, but were exterminated by rats after12 years of predator control (Gaze 2003), possiblya result of the "rat haven" effect (Flux 2002) and thefar faster recovery of rats after control operationswhen other predators are absent. As Elton (1953)concluded, "Cats have a great advantage over humanbeings in the control of rat populations at a very lowlevel: they do not seem to get bored!"

ACKNOWLEDGMENTS

Many thanks to Mike Fitzgerald for his enthusiasticinterest in the diet of our delinquent cat, generous accessto his extensive library on feline ecology, and valuablecomments on early drafts. Felix Manuel Medina mademany constructive comments, and another refereethankfully cut verbiage. Bob Brockie offered comparativematerial from his Wellington cats; CM King edited themanuscript with her usual tact and skill; Meg Flux andother family members helped find kills; and Peng You,the cat, provided all the data.

REFERENCES

Baker PJ, Bentley AJ, Ansell RJ, Harris S 2005. Impact ofpredation by domestic cats Felis catus in an urbanarea. Mammal Review 35: 302-312.

Barratt DG 1997. Predation by house cats, Felis catus (L.),in Canberra, Australia. 1. Prey composition andpreference. Wildlife Research 24: 263-277.

Barratt DG 1998. Predation by house cats, Felis catus (L.),in Canberra, Australia. 11. Factors affecting theamount of prey caught and estimates of the impacton wildlife. Wildlife Research 25: 475-487.

Biben M 1979. Predation and predatory play behaviour ofdomestic cats. Animal Behaviour 27: 81-94.

Borkenhagen P 1978. Von Hauskatzen (Felis sylvestris f.catus L., 1758) eingetragene Beute. Zeitschrift furJagdwissenschaft 24: 27-33.

296 New Zealand Journal of Zoology, 2007, Vol. 34

Brockie B 2001. When cats away the rats will play. TheDominion, Wellington, 22 October 2001.

Bull PC 1953. Aliens in the forest 1. The wild cat problem.Forest & Bird, February, 1953.

Carss DN 1995. Prey brought home by two domesticcats (Felis catus) in northern Scotland. Journal ofZoology (London) 237: 678-686.

Churcher PB 1983. Cats as predators. Unpublished MScthesis in Biology and Education, University ofYork, York.

Churcher PB, Lawton JH 1987. Predation by domesticcats in an English village. Journal of Zoology(London) 212: 439-455.

Courchamp F, Langlais M, Sugihara G 1999. Cats pro-tecting birds: modelling the mesopredator releaseeffect. Journal of Animal Ecology 68: 282-292.

Efford MG, Fitzgerald BM, Karl BJ, Berben PH 2006.Population dynamics of the ship rat Rattus rattusL. in the Orongorongo Valley, New Zealand. NewZealand Journal of Zoology 33: 273-297.

Elton CS 1953. The use of cats in farm rat control. The Brit-ish Journal of Animal Behaviour 1: 151-155.

Finnegan A 1992. Millions of birds may be falling preyto cats. Evening Post, Wellington, 15 October1992.

Fitzgerald BM 1990. Is cat control needed to protecturban wildlife? Environmental Conservation 16:168-169.

Fitzgerald BM, Turner DC 2000. Hunting behaviour ofdomestic cats and their impact on prey popula-tions. In: Turner DC, Bateson P ed. The domesticcat: the biology of its behaviour. Cambridge,Cambridge University Press. Pp. 151-175.

Flux JEC 1997. Status of rabbits (Oryctolagus cuniculus)and hares (Lepus europaeus) in New Zealand.Gibier Faune Sauvage, Game and Wildlife 14:267-280.

Flux JEC 2002. Rat havens. Forest & Bird, May 2002.P. 9.

Flux JEC 2003. Starlings (Sturnus vulgaris) making theirown nest sites. Notornis 50: 238-240.

Flux JEC in press. A review of competition betweenrabbits (Oryctolagus cuniculus) and hares(Lepus europaeus). Proceedings of the SecondWorld Lagomorph Conference, Vairoa, Portugal.Springer Verlag.

Flux JEC, Bull PC 1976. Mammals of the proposedBelmont Regional Hill Park. In: Bagnall RG ed.Survey of the proposed Belmont Regional Park.Wellington, Victoria University. Pp. 61-63.

Flux JEC, Flux MM 1981. Population dynamics andage structure of starlings (Sturnus vulgaris) inNew Zealand. New Zealand Journal of Ecology4: 65-72.

Forest & Bird 2002. Responsible cat ownership. Leafletpublished by The Royal Forest and Bird Protec-tion Society and the Society for the Prevention ofCruelty to Animals. 2 p.

Gaze P 2003. The rise and fall of mohua (Mohoua ochro-cephala) on Mt Stokes. Notornis 50: 176.

George WG 1974. Domestic cats as predators and factorsin winter shortages of raptor prey. Wilson Bulletin86: 384-396.

Gibb JA, Fitzgerald BM 1998. Dynamics of sparse rabbits(Oryctolagus cuniculus), Orongorongo Valley,New Zealand. New Zealand Journal of Zoology25: 231-243.

Gillies C, Clout M 2003. The prey of domestic cats(Felis catus) in two suburbs of Auckland City,New Zealand. Journal of Zoology (London) 259:309-315.

Gillies C, Cutler A 2001. A trial of the liberator, leap-activated, audio visual alarm collar for cats forwildlife protection in Wellington, New Zealand.Unpublished report to The Royal Forest & BirdProtection Society, Wellington, New Zealand.31 p.

Guthrie-Smith H 1953. Tutira. The story of a New Zealandsheep station. Wellington, Reed.

Higgins PJ, Peter JM, Cowling SJ 2006. Handbook ofAustralian, New Zealand and Antarctic birds. Vol.7B. Melbourne, Oxford University Press.

Liberg O 1984. Food habits and prey impact by feraland house-based domestic cats in a rural area insouthern Sweden. Journal of Mammalogy 65:424-432.

Mead CJ 1982. Ringed birds killed by cats. MammalReview 12: 183-186.

Mudge D 2002. Silence of the fantails. New ZealandGeographic 55: 70-86.

Proulx G 1988. Control of urban wildlife predation bycats through public education. EnvironmentalConservation 15: 358-359.

Ruscoe WA, Murphy EC 2005. House mouse. In: KingCM ed. The handbook of New Zealand mammals.2nd ed. Auckland, Oxford University Press. Pp.204-221.

Woods M, McDonald RA, Harris S 2003. Predation ofwildlife by domestic cats Felis catus in GreatBritain. Mammal Review 33: 174-188.