HOME RANGE OF HOUSE CATS FELIS CATUS LIVING WITHIN A NATIONAL PARK

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1 HOME RANGE OF HOUSE CATS FELIS CATUS LIVING WITHIN A NATIONAL PARK PAUL D. MEEK Meek PD, Home range of house cats Felis catus living within a National Park. Australian Mammalogy 25: Fourteen house cats living in residential areas surrounded by National Park were studied using radio telemetry to determine whether they roamed beyond the urban boundary. Eight cats were recorded using natural habitat, predominantly heath the most abundant habitat type adjacent to residential areas. Ninety two percent of fixes were taken within the fringes of the urban boundary. Mean home range size of house cats was 2.9 ha and two categories of cats were identified based on their tendency to wander away from home. Wandering cats had a home range of 5.1 ha and sedentary cats had a range of 0.4 ha. The mean distance travelled by male cats was 70 m and 30 m for females (range m). The longest straight line distance travelled by a house cat in a single foray from a residence was 1.17 km. The major proportion of forays away from the home environs were undertaken at night and in the afternoon. Key words: Domestic cat, home range, radio tracking, activity pattern. P.D. Meek, South-eastern NSW and ACT Hydatid Control Campaign, PO Box Jervis Ba, NSW 2540, Australia. Current address: State Forests of New South Wales, PO Box 535, Coffs Harbour, NSW 2450, Australia. paulme@sf.nsw.gov.au. Manuscript received 26 July 2002; accepted 21 February SEVERAL studies have investigated the ecology of feral cats (Felis catus) in Australia (Jones 1977; Jones and Coman 1981, 1982; Paltridge et al. 1997; Molsher et al. 1999; Molsher 2001) but less is known about house cat ecology. Research on house cat home range, movements, activity patterns, predatory behaviour and impacts was scant in the literature until recent times (Turner and Bateson 2000). In Australia, prey surveys have been undertaken in some states (Paton 1990; Barratt 1997a, 1998; Meek 1998a) and confirm that house cats do kill native wildlife, although the impact on population abundance is unknown (Dickman 1996). Movement studies indicate that domestic cats occupy smaller home ranges than feral cats (Liberg 1984) ranging from approximately 0.02 ha (Bradshaw 1992; Das 1993) to 28 ha (Barratt 1997b). In Cornwall (UK), female farm cats occupied overlapping home ranges between ha (Panaman 1981), and feral cats living on the Avonmouth docks had home ranges of between 10 and 15 ha (Page et al. 1992). A complete overview of the ecology and biology of house cats is presented in Turner and Bateson (2000). The home range and habitat use by house cats with access to resource rich natural environments (abundant native fauna species) have not been comprehensively studied. There is growing pressure in Australia for greater controls over the ownership of house cats (pers. obs.). Numerous local government areas of Australia have introduced, or are in the process of developing legislation to place limits on ownership of pets to make owners more responsible for the behaviour of their pets. In 1991, Victoria (Sherbrooke Council) became the first municipality to introduce a cat management policy (Staindl 1993), and in 1998 the New South Wales parliament passed the Companion Animal Act While research on cats has increased in recent years, some important aspects of domestic house cat biology and ecology is still required to better understand the implications of cats on native wildlife. Dickman (1996) provides a synopsis of our ecological knowledge, however there is still too little scientific information on home range, habitat use, ecological impacts of domestic cats throughout Australia. As such, companion animal legislation/policy is being developed without thorough knowledge of the ecology of domestic house cats. The philosophical position of many agencies and local governments is that cats spread disease to people, foul public areas, invade human space and kill wildlife. In the case of wildlife impacts, too little scientific evidence has been presented to convince pet owners and scientists that house cats do impact on wildlife populations. In Booderee National Park, within the Jervis Bay Territory (a Commonwealth administered enclave on the south coast of New South Wales), the issue of pet

2 52 AUSTRALIAN MAMMALOGY ownership by residents living within the boundaries of the Park was a topic of significant debate during the early 1990 s. Officers of the National Park were concerned that sightings of cats in natural habitat were increasing and there were concerns that house cats were killing native wildlife and posing a threat to threatened species (pers. obs.). The present study was commissioned to provide ecological information to the Jervis Bay Administration for use in the development of companion animal legislation within Jervis Bay Territory. The aims were to collect population data on pet ownership and baseline data on the movements of domestic cats from residential space into the National Park and natural bushland, to: i) determine whether cats roamed in the Park and bushland, ii) determine home range size and maximum foray distances, and iii) highlight the potential ecological threats posed by free-roaming house cats. It is noteworthy that the majority of the cats in this study population were neutered. This paper presents the results of the radio tracking study and discusses management considerations. Details of the prey taken by some of the study cats are discussed in Meek (1998a). METHODS The radio tracking data were collected during a larger study (Meek 1998b) to investigate the movements and biology of foxes (Vulpes vulpes) (Meek and Saunders 2000) and free-roaming dogs (Canis lupus familiaris) (Meek 1999). As such, it mainly includes data collected opportunistically during fox and dog tracking as well as data from specific cat tracking sessions. Site description Bherwerre Peninsula (35 o 13 S, 150 o 45 E) is located 200 km south of Sydney, NSW. It covers approximately 7700 ha and includes Booderee National Park. The peninsula, including the Park and villages are all managed under Commonwealth legislation. The National Park is unique in that three villages are located within its boundaries. Two of these, Jervis Bay Village and HMAS Creswell (RAN) are situated adjacent to each other within an area of 138 ha, bordered on the east by the waters of the Jervis Bay Marine Park with Booderee National Park surrounding the remainder of the residential area (Fig. 1.). The human population of these two villages in 1993/94 was approximately 570 (Australian Bureau of Statistics 2002) and fluctuated due to the recruitment of Navy personnel into HMAS Creswell. The average annual rainfall for Jervis Bay is 1150 mm with May the wettest month and September the driest. Mean maximum day temperature in winter is 15.1 C and 24 C in summer. The area consists of a range of vegetation communities which are dominated by heath (Ingwerson 1976). On the southern edge of the residential area a remnant patch of heath abuts the Park. Booderee National Park is a reserve of high conservation value and represents one of the few remaining intact and unique coastal heath ecosystems on the east coast of Australia (Williams 1995). The Park has high species diversity and wildlife are regularly seen in the residential areas. The fauna include 20 species of extant threatened mammal, bird, amphibian and reptile listed under State and Commonwealth legislation. Choice of cats Study cats were obtained by canvassing cat owners directly and by advertising in the local villages. Each owner was given a brief synopsis of the study and asked if they wanted to volunteer their cats for the study. Cats were selected randomly. All cats were regularly fed by their owners although there were no distinct patterns in feeding times. Cat owners were asked not to curfew their cats during the study to ensure their roaming behaviours were not affected. There are numerous definitions in the literature to describe F. catus in its various forms from wild cats to those of domestic origin (pet). Bradshaw et al. (1999) provides a summary of terms to describe cats depending on their association with man. For the purposes of this study all cats are house cats and later in this paper will be further classified as either wandering or solitary. Radio tracking Radio collars (SIRTRAK, 151 MHz, battery life months) were fitted to cats. Each collar weighed ~35 g. Collars were compact and consisted of a standard leather collar with a resin-covered transmitter package encapsulating reflective tape of different colour combinations. These reflectors aided identification under night lighting. Whip antennae were impregnated with curry and paprika powder to prevent damage by chewing. Radio collars were fitted prior to each tracking session (7 days) and removed by the owners afterwards. A 7-element Yagi antenna attached to a vehicle detected signals that were transmitted into the cabin while driving. Cats were located with a 4-element hand held Yagi antenna. Accuracy of locations was < 20 m as most animals were either observed or known to be close due to the intensity of the signal. Locations (Australian Map Grid coordinates) were recorded and later plotted on maps. Each cat s behaviour and the habitat were recorded at each location. Behaviour was classified when an animal was sighted as either resting, walking, running, grooming, playing, feeding, hunting or marking territory. Habitat was 52

3 MEEK: HOME RANGES OF DOMESTIC CATS 53 Fig. 1. The study site of Jervis Bay Territory, the grey delineates the Booderee National Park. divided into six categories: house and yard, road, heath, forest (coastal eucalypt or tea tree), riparian (creeks and drainage depressions) and grassland (ovals and golf course). Discontinuous tracking (described by Harris et al as random time intervals) was conducted over four nights on each of four occasions (16 nights). Pilot data collection commenced in September 1993 with formal collection between May 1994 and November Opportunistic observations were taken at other times throughout the study. Twelve, 18 and 24 hr sampling sessions were conducted which enabled collection of data during the daylight hours to ensure both diurnal and nocturnal fixes were included in the analysis. Fixes on study animals were attempted every hour throughout the sampling periods commencing on the next nearest hour since the last observation. Barratt (1997b) found that one hour intervals for suburban cats was sufficient to overcome the problem of statistically independent data (White and Garrott 1990). Home range analysis Animal location data were digitised using a Graphical Facilities Information System (AUTOCAD-VQ) based on the Australian Map Grid (zone 56). Accuracy of the AUTOCAD program was checked by ground-truthing the distance between known points using a Kaycee circular measuring wheel. The accuracy of the package was < 3 m on all field tests. Home range area was calculated using CALHOME (Kie et al. 1994) and HOMERANGE (Ackerman et al. 1990). The Minimum Convex Polygon (MCP) (Southwood 1966) and Adaptive Kernel methods were used to determine home range. The Adaptive Kernel method was not suitable for analysing some of the data as it was unable to deal with the clumped distribution/duplication of records within a household, regardless of manipulation of the smoothing value (h) as described by Worton (1989). Consideration was given to removing all records in and around the household to remove their clumped distribution, however this would have further biased the data and excluded many cat home ranges from the study. Given that identifying movements of cats outside of the household environment was a critical component of the study, it was decided to calculate home ranges using MCP 100% confidence limits. Using 95% confidence limits removes outliers therefore excluding forays and infrequently visited sites (White and Garrott 1990). In this study, outliers were important and removing them would have significantly reduced the home range size for some cats thereby defeating the purpose of the investigation. Mean distance travelled was calculated by the home range package between each location recorded and do not reflect consecutive movements between locations during a foray. Home range data were mapped using a geographical information system (ARCVIEW3) to produce visual displays of range size and shape. 53

4 54 AUSTRALIAN MAMMALOGY RESULTS Home range and movements Although 20 domestic cats were radio collared, location data from only 15 were analysed because tracking effort was interrupted when some cats owners moved out of the residential area. Overall, 2202 locations were collected across the study population and the number of fixes on cats ranged between 48 and 356 (Table 1). Thirteen study cats (65%) were de-sexed, five were entire and the status of two was undetermined. The sex ratio of the study cat population was 2F:1M (13 females and 7 males) (Table 1). The mean home range (+SE) of house cats in Jervis Bay as defined by minimum convex polygon (100%) was ha, the median value was 1.5 ha (Table 2). Male cat mean home range size was ha and female home range size was ha. These were not significantly different (t = - 0.7, df = 13, P = 0.5). Following Bradshaw et al. (1990) and Das (1993) wanderers in this study are defined as cats that roamed outside of their household domain (> 1 ha). Seven of the 15 study cats were classed as wanderers' based on the above description and a significant difference in home range size (t = - 2.6, df = 7.1, P = 0.03). The mean home range (+SE) of wandering cats was ha (median 2.8 ha), the sedentary cats had home ranges of ha (median 0.4 ha) (Table 2). F1, a desexed cat had the largest home range recorded (15.65 ha: Table 1). This animal was often recorded (24% of fixes) hunting rabbits (Oryctolagus cuninculus) and rats (Rattus rattus) behind the residential area of Jervis Bay Village in heath. M1, also desexed, resided in the same house and had a large home range (12.04 ha: Table 1) of which 25% of fixes were in heath. Home ranges of all radio collared house cats from within and between residences were spatially (Fig. 2) but not temporally overlapping. Fighting between cats was observed on a few occasions on the roads between residences. There was no difference in home range size between entire and desexed animals (t = -0.7, df = 13, P = 0.5) although the sample size of entire animals was very small (N = 2). The maximum linear distance travelled by an individual cat was 1170 m in one night (M7). This cat also travelled the largest mean distance per hour (272 m) although it only had four fixes. F1 was regularly recorded hunting 370 m from home. F13 was trapped on the boundary fence line to Jervis Bay Range Facility (JB Airfield) early in the study, approximately 100 m from its residence (Fig. 1). The population mean calculated from the individual mean distance as determined by the home range package (+ Name ID Sex Status Breed Age (yr) N MCP (ha) Mean distance travelled 100% 95% (m) Fluff F1 F D Tabby Orange Boy M1 M D Ginger Tinkerbell F2 F D Tabby Sarg M2 M D Tortoise shell Cassie F3 F D White and red Candy F4 F D Ginger Poncho M3 M E Persian x tabby Sampson M4 M D Persian Emma F5 F D Tortoise shell Tiffiny F6 F D Tortoise shell Chloe F7 F E Tabby Katie F8 F D Grey and white Peggy F9 F D Cream and white Puss F10 F D Persian Sadie F11 F D Tabby Moggie* F12 F U Black and white Sam* M5 M U Orange x Tabby Boof* F13 F E Ginger/grey Ginger* M6 M E Ginger Frank* M7 M E Black Table 1. Data on home range (MCP) and mean distance travelled for 15 cats from Jervis Bay over N is number of radio fixes. * denotes that a cat left the study before enough data could be collected. D = desexed, E = entire, F = female, M = male and U = unknown status. 54

5 MEEK: HOME RANGES OF DOMESTIC CATS 55 Number of cats Mean home range (ha + SE) Range (ha) Mean Distance travelled (m + SE) Wandering Sedentary Table 2. Home range and mean distance travelled of wandering and sedentary house cats in Jervis Bay. SE) for male cats (N = 5) was m and females (N = 12) m. These values were not significantly different (t = 1.1, df = 15, P = 0.3). Activity observations There were two spikes in foray activity (Fig. 3), one at night (2400 hrs) and in the late morning (1100 hrs). The lowest activity hour was M1 was regularly observed resting in the roadside verge of heath adjacent to his residence during daylight hours. It is difficult to categorically determine the proportion of visits that were undertaken to hunt rodents compared to forays to resting sites, although M1 did bring R. rattus, Antechinus spp. and house mice (Mus domesticus) home after nocturnal forays (Meek 1998a). Habitat use Forty six percent of the population were recorded in natural habitat at some time during the tracking study, although the frequency of occurrence of cats in natural habitat was low. Most fixes were recorded within the urban environment (91%), either in the cats home and yard or on the streets. The remainder of records were in natural habitat, predominantly heath (8%) (Table 3). Wandering cat fixes were mostly in urban environs, however 12% of fixes were in heath. Forest, ovals and riparian habitat were used, albeit infrequently. DISCUSSION The size of home ranges estimated in this study for house cats in Jervis Bay are similar to those reported by other researchers (Panaman 1981; Page et al. 1992; Das 1993; Barratt 1997b) and all fall within the ranges ( ha) reported by Panaman (1981) at the 95% contour level. The home ranges of seven cats included natural bushland in some form and there was spatial overlap in home ranges. Barratt (1997b) found that cats living in the same house had either completely overlapping home ranges and/or core areas depending on their relatedness to one another. In this study, the home ranges of cats living in the same house and in different houses did overlap extensively although not at the core activity level (household area), with the exception of those cats in the same residence. There was an obvious dichotomy in behaviour between groups of cats and I have referred to them as wandering and sedentary cats. The reasons why some cats roamed and others were sedentary could not be determined, although on a few occasions some cats were kept at home during the night. This occurred infrequently for two cats and would not have biased the results as the data was not collected when it was clear the cats were not freeroaming. Owners were asked not to lock animals in doors during data collection and mostly honoured this request. Konecny (1987) found that patchily distributed food led feral cats to have large home ranges. The same may have occurred in this study because some cats displayed a preference for certain prey types and these were not always close to their residence. These cats had preferred hunting sites that were mostly visited at night. Barratt (1997b) also found that house cats in Canberra had favoured hunting sites and used familiar travel routes to access these sites. Liberg et al. (2000) believed that home range of male cats in their study was influenced by access to females for reproduction. The same influences do not seem to be affecting the cats in this study although this result is biased by the number of females and de-sexed males; the two largest home ranges were a de-sexed male and female from the same household. Barratt (1997b) reported similar home range sizes in three suburban cats (1 entire and 2 desexed) and agreed with other authors that home range size was influenced by food resources, kinship and distribution of females. There was no difference in home range size between male and female cats in this study although there were only two entire cats in the study population and it is not known what effect neutering has on territoriality and roaming behaviour. Summer activity patterns of feral cats suggest a crepuscular mode, although this pattern changes during winter with increased diurnal activity (Izawa 1983). Cats in this study displayed similar behaviours to house cats in Canberra (Barratt 1997b) in that hunting forays were nocturnal, although a second spike in activity occurred in the afternoon when cats sought resting sites. Interestingly, the cats seemed to be less active around hrs which could be related to the time when most families return from work. When the cats were on a hunting foray it was noticeable that they did not travel in open country and walked close to fence lines and vegetation boundaries where they were not easily detectable. Roads and tracks were used as navigational paths but no cat was seen walking down a road during a known 55

6 56 AUSTRALIAN MAMMALOGY Fig. 2. Overlapping home ranges of six cats from the Jervis Bay village Percentage of Fixes Time (24 hr) Fig. 3. Percentage of fixes of radio collared cats outside of the home environs (all cats combined) for each EST hour. 56

7 MEEK: HOME RANGES OF DOMESTIC CATS 57 Cats House / yard Heath Road Forest Grassland Riparian Wandering N Mean percentile <1 <1 <1 Sedentary N Mean percentile 94 <1 5 - <1 - All cats N Mean percentile <1 <1 <1 Table 3. Mean percentage of fixes of radio collared cats in various habitat types. N = number of radio fixes per habitat type. hunting foray. Cats were not seen roaming together at any time during the study, although radio tagged cats were often in close proximity to radio tagged foxes. Some of the cats (M3, F11, F6, F4, F3) showed high site fidelity to one or two houses (reflected in the radio tracking data). Cats from separate houses showed avoidance of one another, although home ranges did spatially over-lap. The number of fixes in natural bushland was considerably fewer than in urban environs. However, the information in this study and data presented in Meek (1998a) show that wildlife interactions with house cats are frequent given the proximity of residential areas to food and shelter resources of natural bushland. Several threatened mammal species are known to have been hunted by cats in Jervis Bay (Meek 1998a; Meek and Triggs 1998a; Meek and Triggs 1998b) and several threatened birds including the ground parrot (Pezoporus wallicus) and eastern bristlebird (Dasyornis brachypterus) are at risk of predation in this area. Known records of the eastern chestnut mouse (Pseudomys gracilicaudatus) and common dunnart (Sminthopsis leucopus) have been found in close proximity to residential areas, the latter having been reported in a cat scat (Meek and Triggs 1998a). Freeroaming cats close to natural bushland where densities of threatened species are high may have a deleterious effect on prey population abundance. This edge effect occurred in England where the number of prey caught was higher for cats living on the fringes than those in the town (Churcher and Lawton 1987). Likewise, Trueman (1990) found house cats living close to natural bushland caught more birds than those in urban areas. Cats in this study were observed stalking, catching and killing native wildlife and several cat owners described their pets bringing home badly injured wildlife (Meek 1998a). On one occasion the author was called to a household to remove an adult sugar glider (Petaurus breviceps) that had its chest cavity torn open and was still alive under the owners bed with young attached to its body. Fitzgerald and Turner (2000) suggest it could be a phenomenon of domestication that cats are returning the hunted game to their owner. Leyhausen (1979) believed that the behaviour of returning home with prey may be related to parental training where the owner is a deputy kitten. Limits on roaming behaviour in kittens may reduce their preponderance to hunt. At the onset of this study all cat owners claimed that their cats never roamed and rarely killed any wildlife apart from a few mice all were completely amazed when they saw evidence of their cats home range. Early in the study two cats from HMAS Creswell were trapped 2.5 km from their house. All other cats went on forays of up to 600 m from their houses. In areas where threatened species are in close proximity to residential areas, these forays may pose a significant threat, particularly where the resident cat has a predilection for hunting. Interestingly, when cat home ranges were plotted on a land tenure map, none of them extended into the formal National Park boundaries, although there was contiguous habitat with HMAS Creswell and Jervis Bay village that was used by the cats. Observations of hunting behaviour during tracking varied between cats and was also reflected in prey data collected on this population (Meek 1998a). It was apparent that certain cats showed a particular interest or ability to hunt specific wildlife as reported by Barratt (1997b; 1998). F2 seemed to prefer hunting P. breviceps and ring-tailed possums (Pseudocheirus peregrinus) while F1 and M1 regularly caught bush rats (Rattus fuscipes), O. cuniculus, R. rattus and M. domesticus. This individual variation in cat behaviour has also been discussed in detail by McCune (1995) and Mendl and Harcourt (2000). The management of cats in natural habitat areas should consider a prohibition on free-roaming. Caro (1980) found that kittens learned to hunt and kill prey from their mothers within the first eight weeks of life. As such, continuous curfews on domestic cats may reduce exposure to prey and may in time (generations) modify cat behaviour away from hunting prey. Curfews also place more emphasis on close contact between adult cats, humans and the kittens during the early months of life which may 57

8 58 AUSTRALIAN MAMMALOGY increase the kittens long term friendliness to people (McCune 1995). The results of a pet survey in Jervis Bay (Meek 1994) reported that pet owners represented 71% of the Jervis Bay Territory population. Cats were kept by 26% of households. At the time of this study there was no formal pet registration, so cats could roam and breed freely. Registration of pets often puts the responsibility of ownership directly on the owner (Murray and Penridge 1992) which, in the long-term, means benefits for the pet and the surrounding environment. The introduction of cat registration, prevention of free-roaming behaviour and compulsory de-sexing in local government areas will contribute to reductions in the number of unwanted cats and lead to more responsible ownership which ultimately reduces the potential impact on native species. However, it should be noted that data collected in this study show that de-sexed cats will still roam and hunt wildlife. As such de-sexing should not be seen solely as a mechanism to reduce hunting and roaming behaviour. To date the Jervis Bay Territory does not have companion animal legislation and the information presented here indicates that if regulations were introduced that specify desexing and curfews of cats it still wont adequately modify cat behaviour. There is a growing body of evidence to confirm that some of the management strategies believed to reduce cat predation are inappropriate. Trueman (1990) found that cats with bells on their collars do not catch less prey than un-collared cats. Biben (1979) used experiments to determine that hunger was not a driving factor in killing behaviour by cats. Barratt (1997b) and this study suggest that regular feeding by owners will not reduce the cats need to hunt and kill prey (also see Fitzgerald and Turner (2000). The findings of this study support the research of Barratt (1997a,b) that house cats living in close proximity to nature reserves with abundant wildlife may pose a risk to threatened species. The unresolved question is whether cat predation is affecting species population dynamics. In the villages of Jervis Bay and HMAS Creswell, natural bushland is very close to residential areas and all native species are at risk of predation by some of the house cat population. Hunting behaviour modification through appropriate cat management legislation is yet to be tested in practice, and even if behaviours are modified by better ownership practices, we still need to know what impact cats can have on native wildlife where other forces (eg. fox predation and habitat fragmentation) are also contributing to species declines. ACKNOWLEDGMENTS Research was conducted under Booderee National Park permit number JBLA 22. Animal ethics approval was granted by the University of Canberra Animal Experimentation and Ethics Committee (AE 93/5). Funding was provided by the Wildlife, Exotic Disease Preparedness Program, Bureau of Rural Resources and Jervis Bay Administration. Thanks to Peter Lawler and Linda Hodges for supporting the research program. Richard Hawksby, Athol Ardler, Yvonne Brown, Ursula Moore, Teresa Thies, Rachael Dimasi, Vivina Sinnamon and Phil Borchard provided field assistance. Thank you to the cat owners of Jervis Bay Village and HMAS Creswell. Nick Dexter and Sally Radford made helpful comments on this manuscript. Thanks to Sally Radford for sharing her computing skills and for having faith in my data. REFERENCES ACKERMAN B, LEBA F, SAMUEL M AND GARTON E, User s manual for program home range. Idaho Forest and Wildlife Range Experiment Report 15: AUSTRALIAN BUREAU OF STATISTICS, Census data BARRATT DG, 1997a. Predation by house cats Felis catus (L.), in Canberra. I. Prey composition and preference. Wildlife Research 24: BARRATT DG, 1997b. Home range size, habitat utilisation and movement patterns of suburban and farm cats Felis catus. Ecography 20: BARRATT DG, Predation by house cats, Felis catus (L.), in Canberra, Australia. II. Factors affecting the amount of prey caught and estimates of the impact on wildlife. Wildlife Research 25: BIBEN M, Predation and predatory play behaviour of domestic cats. Animal Behaviour 27: BRADSHAW JWS, The behaviour of the domestic cat. CAB International: Wallingford. BRADSHAW JWS, HORSFIELD GF, ALLEN JA AND ROBINSON IH, Feral cats: their role in the population dynamics of Felis catus. Applied Animal Behaviour Science 65: CARO TM, Predatory behaviour in domestic cat mothers. Behaviour 74:

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