Research Studies on the Behavior of

Outdoor-Access and Free-Ranging Domestic Cats

Bibliography with Abstracts
Compiled by Danielle Robertson
Compassionate Pet Services
P.O. Box 801, Granby, MA 01033

Baker, Philip J., Carl. D. Soulsbury, Graziella Iossa, and Stephen Harris. 2010. Domestic
Cat (Felis catus) and Domestic Dog (Canis familiaris) in Urban Carnivores: Ecology,
Conflict, and Conservation. Gehrt, Stanley, D., Seth P.D. Riley, and Brian L. Cypher,
editors. The John Hopkins University Press.

Barrat, David. 1997. Home range size, habitat utilisation and movement patterns of
suburban and farm cats Felis catus. Ecography 20(3): 271–280.
Abstract: The movements of 10 house cats (4 desexed females, 5 desexed males and 1 intact
male) living on the edge of a suburb adjoining grassland and forest/woodland habitat, and a
neighbouring colony of seven farm cats, were examined using radio-telemetry over nine months.
Nocturnal home range areas of the suburban cats varied between 0.02 and 27.93 ha (mean 7.89
ha), and were larger than diurnal home range areas (range 0.02 to 17.19 ha - mean 2.73 ha).
Nocturnal home range areas of cats from the farm cat colony varied between 1.38 and 4.46 ha
(mean 2.54 ha), and were also larger than diurnal home range areas (range 0.77 to 3.70 ha - mean
1.70 ha). Home ranges of cats in the farm cat colony overlapped extensively, as did those of cats
living at the same suburban residence. There was no overlap of home ranges of female cats from
different residences, and little overlap between males and females from different residences.
Four of the suburban house cats moved between 390 m and 900 m into habitat adjoining the
suburb. Polygons describing the home ranges of these animals were strongly spatially biased
away from the suburban environment, though the cats spent the majority of their time within the
bounds of the suburb. Movements further than 100-200 m beyond the suburb edge were always
made at night. There is evidence that home range sizes and spatial movement patterns of house
cats are largely determined by: a) the density and spatial distribution of cats utilizing separate
food resources, b) the personality and social dominance of individual cats, c) the location of
favoured hunting and resting/sunning sites, and, d) barriers such as busy roads.

Calhoon, R.E. and Haspel. 1989. Urban cat populations composed by season, subhabitat
and supplemental feeding. Journal of Animal Ecology 58: 321-328.
Abstract: (1) Population densities of free-ranging cats were compared in two contiguous
urban subhabitats, in three seasons, and in response to supplemental feeding. (2) One subhabitat,
characterized by voluminous, poorly contained refuse, and many abandoned buildings, supported
4.88 +-0.82 cats ha-1 (mean +-S.D.), which differed significantly from the 2.03 +-0.2 cats ha-1
supported by the other subhabitat (partial refuse containment, few abandoned buildings). (3)
Neither season nor supplemental feeding had a significant effect on population density. (4) The
distribution of individuals within the study area varied with the availability of shelter and was
not dependent upon food.
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Childs, J.E. and Ross L. 1986. Urban cats: characteristics and estimation of mortality due
to motor vehicles. American Journal of Veterinary Research 47(7): 1643-8.
Abstract. The present study was undertaken to characterize and estimate cat mortality caused by
motor vehicles in Baltimore, MD. Analysis of Municipal Animal Shelter records revealed that
the number of dead cats annually removed from Baltimore streets averaged 2,721 over 3 years.
The number of cats removed per year was notably constant (range 2,621 to 2,917), with a
consistently greater number of animals being picked up during summer months. A map of the
location of 634 dead cats removed from city streets showed that areas of highest cat mortality
were associated with areas of highest human population density. This map was used to stratify
Baltimore into areas of high, medium, and low cat mortality for subsequent stratified random
placement of tagged dead cats for the mark-recapture study. One hundred fifteen tagged dead
cats were placed on city streets during the course of 1 year, and 50.5% were retrieved by the
Municipal Animal Shelter. We estimate over 5,000 free-ranging cats are killed by
automobiles/year in Baltimore. Analysis of 212 dead cats removed from city streets showed that
the majority were male (63%) and that most animals were sexually intact (90%). At least 20% of
the dead animals was previously owned, and few kittens or juvenile cats were found in the
sample.

Devillard, Sebastien, L. Say, and D. Pontier. 2003. Dispersal pattern of domestic cats (Felis
catus) in a promiscuous urban population: do females disperse or die? Journal of
Animal Ecology 72: 203-211.
Abstract: 1. The domestic feral cat (Felis catus L.) is a good model for studying intraspecific
variability of dispersal patterns in mammals because cats live under a large diversity of socio-
ecological conditions. We analysed both the natal and breeding dispersal patterns of domestic
cats in a promiscuous urban population and tested whether or not it differed from the male-
biased natal dispersal pattern observed for polygynous rural populations. 2. During an 8-year
study we recorded the exact date of in situ death for 148 marked cats and the exact date of
disappearance from the population for 99 other cats. Because undiscovered deaths might over-
estimate dispersal probabilities when considering only disappearance probabilities, we made an
novel application of multistrata capture-recapture methods in order to disentangle dispersal from
true mortality. 3. We showed that mature females dispersed, both before and after their first
reproduction, at 1 and 2 years old. Contrary to females, no dispersal seemed to occur in males.
Before sexual maturity, females that disappeared at 1 and 2 years old were in worse body
condition than females that stayed in the population area after 2 years old. However, they did not
reproduce less successfully before their disappearance than females that died later in the
population area. 4. The female-biased and low natal dispersal pattern in this population was
atypical compared to other promiscuous/polygynous mammals and differed from that observed
in rural polygynous populations of domestic cat. Neither local mate competition nor inbreeding
avoidance appeared to be sufficient pressures to counterbalance ecological constraints on
dispereal in an urban environment. However, local resource competition for den sites between
potential matriarchies could lead to the breeding dispersal of less competitive females.

Germain, E., S. Benhamou 3 & M.-L. Poulle . 2008. Spatio-temporal sharing between the
European wildcat, the domestic cat and their hybrids. Journal of Zoology 276(2):
195-203.
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Abstract: The European wildcat Felis silvestris silvestris, which can hybridize with the domestic
cat Felis catus to produce fertile hybrids, is threatened by hybridization. To identify the
behavioural processes that can affect interbreeding, we investigated the spatio-temporal sharing
between wildcats, domestic cats and their hybrids (defined on their genotypes) in a rural area of
north-eastern France where hybridization is frequent. Wildcats' and hybrids' home ranges were
larger than those of domestic cats, and they did not vary according to body mass, season and
reproductive period. The three types of cats had similar daily activity rhythms but the
concordance between their space use patterns was low or null. Thus, a high spatio-temporal
concordance is not a prerequisite for hybridization. Rare excursions made by the cats outside of
their home ranges may be at the origin of interbreeding. Moreover, hybrids may play a key role
in hybridization by behaving as wildcats and by sharing at least a part of their range with them as
well as with domestic cats. Behavioural barriers between them and wildcats may not exist
because of their similarity in morphology and spatial behavior.

Goszczynski, J., D. Krauze, and J. Gvyz. 2009. Activity and exploration of house cats in
rural areas of central Poland. Folia Zoologica 58(4): 363-71.
Abstract. Domestic cats are the most numerous predators in Poland. They are commonly kept
at farms but hardly controlled, so penetrate freely wide range of habitats. The work aimed at
determining the range of greatest impact of cats by identifying patterns of their activity and area
searching, over daily, monthly and annual cycles. The density index, estimated from transect
counts, performed along standard routes, proved to be dependent on temperature, precipitation
and time of the day. In spring and summer, cats presented a two-peaked activity pattern, while in
cold seasons it was more stable throughout the day. In warm months cats were registered at a
further distance from the buildings than in colder ones. The animals were much less active when
rain was falling. Cats' responses on noticing an observer showed that the further they were from
the edges of settlements the more timid and cautious they became. The results showed that the
potential pressure that cats may exert on their prey is the biggest around dawn and dusk and in
summer. During a daytime it is confined to the immediate vicinity of build-up areas.

Grubbs, Shannon E. and Paul R. Krausman. 2009. Observations of Coyote – Cat

Interactions. Journal of Wildlife Management 73(5): 683-685.

Harper, Grant A. 2007. Habitat selection of feral cats (Felis catus) on a temperate, forested
island. Austral Ecology 32: 305-314.
Abstract. Habitat selection of mammalian predators is known to be influenced by availability
and distribution of prey. The habitat selection of feral cats on Stewart Island, southern New
Zealand, was investigated using telemetry of radio-tagged cats. Compositional analysis of the
habitat selection of radio-tagged cats showed they were using the available habitats non-
randomly. Feral cats avoided subalpine shrubland and preferentially selected podocarp-broadleaf
forest. The avoidance of subalpine shrubland by cats was probably due to a combination of the
presence of a large aggressive prey species, Norway rats Rattus norvegicus, and the lack of rain-
impervious shelter there. Most cats also used subalpine shrubland more often in dry weather than
in wet weather. Cats did not preferentially select all the other habitats with only smaller rat
species, Rattus rattus and Rattus exulans, present however. Cats were probably further
influenced by the availability of large trees, in podocarp-broadleaf forest, that can provide
shelter. Cats were also more active in dry rather than wet weather which supports this
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conclusion. Home ranges of feral cats on Stewart Island were some of the largest recorded,
probably because of limited primary and alternative prey.

Haspel, Carol and Robert Calhoon. 1993. Activity patterns of free-ranging cats in Brooklyn,
New York. Journal of Mammalogy 74(1): 1-8.
Abstract: Activity patterns of free-ranging cats in Brooklyn, New York, varied with
neighborhood, season, and hour of the night. Standardized partial-regression analysis identified
temperature, humidity, precipitation, and hour as accounting for 32.6% of feral-cat activity. Cats
in the area of private homes included free-ranging pets while those in the area of multiple
dwellings were commonly feral. Regardless of neighborhood, nighttime activity was bimodal
with peaks occurring at 0100 h and at sunrise. Activity levels declined throughout the autumn
and increased in the spring.

Haspel, C. and R. Calhoon. 1989. Home ranges of free-ranging cats (Felis catus) in
Brooklyn, New York. Canadian Journal of Zoology 67(1): 178-181.

Kays, Roland and Amielle DeWan. 2004. Ecological impact of inside/outside cats around a
suburban nature preserve. Animal Conservation 7: 1-11.
(Online: http://www.nysm.nysed.gov/staffpubs/docs/15128.pdf )
Abstract: While subsidised populations of feral cats are known to impact their prey
populations, little is known about the ecological impact of inside/outside hunting cats (IOHC).
We studied IOHC around a suburban nature preserve. Mail surveys indicated an average of 0.275
OHC/house, leading to a regional density estimate of 0.32 IOHC/ha. A geographical model of
cat density was created based on local house density and distance from forest/neighbourhood
edge. IOHC hunted mostly small mammals, averaging 1.67 prey brought home/cat/month and a
kill rate of 13%. Predation rates based on kills brought home was lower than the estimate from
observing hunting cats (5.54 kills/cat/month). IOHC spent most outside time in their or their
immediate neighbours’ garden/yard, or in the nearby forest edge; 80% of observed hunts
occurred in a garden/yard or in the first 10m of forest. Radio-tracked IOHC averaged 0.24 ha in
home range size (95% minimum convex polygon (MCP)) and rarely entered forest. Confirming
this, scent stations detected cats more often near the edge and more cats were detected in smaller
forest fragments. There was no relationship between the number of cats detected in an area and
the local small mammal abundance or rodent seed predation rates. Cold weather and healthy cat
predator populations are speculated to minimise the ecological impact of IOHC on this area.

Langham, N.P.E. and R.E.R. Porter. 1991. Feral Cats (Felis catus L.) on New Zealand
Farmland. I. Home Range. Wildlife Research 18: 741-760.
Abstract. The movements of a population of feral cats (Felis catus) were monitored on New
Zealand farmland over three years by means of radiotelemetry. The number of resident males on
the 5.2 km2 study area varied from 5 to 9, averaging 1.34 per km2, compared with 10-13 females,
averaging 2.19 per km2. The average against the over three years was 3.47 cats per km2.
The nocturnal home range was significantly larger than the diurnal home range in both sexes.
Adult females home ranges overlapped considerably; adult males tended to occupy exclusive
home ranges or territories with little overlap, but including those of several females. Adult males

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and females that used barns as den sites were mainly nocturnal and had larger home ranges than
females denning in vegetation.
Females showed no consistent changes in home range size with season, although those breeding
in barns had larger home ranges in summer. Adult males had larger territories in summer and
winter. Dispersing subadult males had a similar home range to adult males. Death or
disappearance of a dominant male allowed new males to occupy the vacated territory. Two
subadults were tracked by day until they became adult and acquired territories within the study
area.

Langham, N.P.E. 1992. Feral Cats (Felis catus L.) on New Zealand Farmland. II. Seasonal
Activity. Wildlife Research 19: 707-720.
Abstract. The activity patterns of a resident population of 15 feral cats (Felis catus) on New
Zealand farmland were investigated from March 1984 until February 1987 by radiotelemetry.
Females could be divided into two separate groups: (1) those denning in barns and (2) those
denning in the swamp and willows. Females denning in barns were mainly nocturnal except in
spring and summer when rearing kittens. Barn cats moved significantly further between dusk
and dawn, except in autumn-winter, than those denning in swamp and willows which were active
over 24 hours. When not breeding, related females occupied the same barn. In both groups, the
home range of female relatives overlapped. Males ranged over all habitats, and dominant males
moved significantly further and had larger home ranges than other males in all seasons, except in
summer when they rested, avoiding hot summer days. Only adult males were active during the
day in spring and autumn-winter. The importance Zeitgeber in synchronizing cat activity with
that of the prey is examined. The significance of the female den site is discussed in relation to
proximity of food, predators, and social behavior and male defense.

Liberg, O., M. Sandell, D. Pontier, and E. Natoli. 2000. Spatial organization and
reproductive tactics in the domestic cat and other felids. Pages 120-147 in Turner, D.
C., and P. Bateson, eds. The domestic cat: the biology of its behavior. Cambridge Univ.
Press, New York, NY.

Liberg. O. 1984. Home range and territoriality in free ranging house cats. Acta Zoologica
Fennica 171: 283-185.

Liberg, O. 1980. Spacing patterns in a population of rural free roaming domestic cats.
Oikos 35: 336-349.
Abstract. Spacing patterns in a population of domestic and feral cats in a rural area in southern
Sweden were investigated by visual observations, trapping and radio tracking. Females lived
alone or in groups around human households. Within each female group the home ranges almost
completely overlapped, but between different female groups there was little or no overlap. Most
females remained in the same place all their lives, but a few individuals moved and became
established at new households, invariably one where there were no other female cats. There were
always six to eight feral, well established males in the area, with moderately overlapping home
ranges. These ranges were considerably larger than those of females, and one male might include
several female groups within his home range. Young males, born in the area, stayed with the
female group, where they were born until they were 1.5-3 yr old. They then left and tried to settle
somewhere else. Spacing patterns in this cat population can be explained by the influence of
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proximate and ultimate factors, among which intraspecific aggression and adaptation to living in
human households are the most important. Parallel evolution of lion and house cat social
organizations is discussed.

Marks, Britni K. and R. Scot Duncan. 2009. Use of forest edges by free-ranging cats and
dogs in an urban forest fragment. Southeastern Naturalist 8(3): 427-436.
Abstract. Free-ranging Felis catus (Domestic Cat) and Canis familiaris (Domestic Dog) can
greatly impact native prey populations, but little is known about their occurrence in urban forest
fragments. In this study, we used camera traps to photograph (capture) cats, dogs, and native
wildlife in a 409-ha urban forest in Birmingham, AL from Jan–Apr 2007. Habitat treatments
included forest interior and forest edges by industrial lands, neighborhoods with higher house
values, and neighborhoods with lower house values. We employed both conservative (n = 31)
and liberal (n = 64) methods of tallying the number of individual dogs, cats, and native mammals
captured. Dogs and cats combined comprised 19% (conservative) and 26% (liberal) of all
photographic captures. Procyon lotor (Raccoon) were the most abundant of the 7 native species
at 32% (conservative) and 53% (liberal) of all captures. Dogs were more abundant in
neighborhood edges, and cats were more abundant in the forest interior. Cats and dogs combined
were 75% (conservative) and 86% (liberal) of captures from the forest interior. Captures of
native species were far more frequent in neighborhood edges (conservative = 86.9%, and liberal
= 92.3%) than in other treatments. These findings demonstrate that exotic predators can be an
important ecological presence in certain portions of urban forest fragments, and more extensive
studies of their impact are needed.

Meek, Paul. 2003. Home range of house cats Felis catus living within a National Park.
Australian Mammology 25: 51-60.
(Online: http://www.publish.csiro.au/?act=view_file&file_id=AM03051.pdf )
Abstract: 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 1.5 – 272 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.

Molsher, Robyn, C. Dickman, A. Newsome, and W. Muller. 2005. Home ranges of feral cats
(Felis catus) in central-western New South Wales, Australia. Wildlife Research 32:
587-595.
Abstract: Twenty-one feral cats were radio-tracked using direct sighting and triangulation
techniques (amassing 730 location fixes) during winter in an agricultural landscape in central-
western New South Wales. Factors affecting home-range size, home-range overlap and habitat
use were assessed. Mean home-range size was 248 ha (s.e. = 34.9, n = 15 cats, 598 location
fixes). Home-range size and habitat use were not influenced by sex or age of adult cats, prey
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abundance or time of day. However, cat weight significantly influenced range size, with heavier
cats having larger ranges than smaller cats. Although the cats are apparently solitary, their home
ranges overlapped considerably, particularly between young adults and old adult cats. Cats were
active both by day and night and did not occupy permanent dens. Home ranges encompassed
mixed habitat types that provided both shelter and prey. Open woodland and open forest were the
main habitat types covered by home ranges, but within these areas cats showed a preference for
grassland, where rabbits were more abundant. The results recorded in this study indicate that cat-
control programs should concentrate in mixed habitat areas, where both shelter and food are
available, and over widely dispersed areas. The absence of group living suggests that the
effectiveness of virally vectored fertility or biological control agents would be limited.

Morgan, S.A., C.M. Hansen, J.G. Ross, G.J. Hickling, S.C. Ogilvie, and A.M. Paterson. 2009.
Urban cat (Felis catus) movement and predation activity associated with a wetland
reserve in New Zealand. Wildlife Research 36: 574-580.
Abstract: Context. House cats are increasingly suggested as having major ecological impacts
in semiurban environments. Information on the activity of house cats is relatively scarce,
especially in habitats such as wetlands. Aims. This study examines the movement and foraging
behaviour of house cats living on the periphery of a wetland reserve in Christchurch city, New
Zealand. Methods. Twenty-one domestic cats living in a suburban residential area were studied
using radiotelemetry to determine home-range size, mean and maximum distances travelled into
the adjacent wetland, and the proportion of time spent in the wetland over a 12-month period.
Surveys of prey retrieval for 88 cats were also carried out by cat owners over the same 12-month
period. Key results. Cat age and the distance of the cat’s home from the periphery of the wetland
were highly correlated with cat movement and hunting activity. These movements were not
markedly influenced by season or time of day. Younger cats (<6 years of age) living on the
periphery of the wetland had larger home-range sizes, moved significantly further into the
wetland and spent a significantly greater proportion of time in the wetland. Cats living close to
the wetland also brought a greater diversity and a greater total number of prey items to their
home-site. Rates of predation were not significantly influenced by sex or whether the cat was
wearing a bell. The most common prey items were introduced rodents and birds; however, 172
of 981 prey items were identified as a native common skink. Conclusions.Consequently, cats
living in households on the wetland periphery currently pose a predation risk for the wetland
species, and the impact of cats on the native skink population warrants further investigation.
Implications. This study suggests that domestic cats will exploit wild habitats but that their
potential impact will have both positive (predation of introduced pest species) and negative
(occasional direct predation) effects on native wildlife.

Ogan, Chester and Jurek Ronald. Biology and ecology of feral, free-roaming and stray
cats. Pages 87-91 in Mesocarnivores of Northern California: biology, management &
survey techniques.
(Online: http://www.fs.fed.us/psw/publications/4251/ogan1.PDF )

Olaf, Liberg. 1984. Food habits and prey impact by feral and house-based domestic cats in
a rural area in southern Sweden. Journal of Mammalogy 65(3): 424-432.
Abstract. Natural prey of domestic cats (Felis catus) in the Revinge area in southern Sweden
during 1974-79 was related to prey abundance, annual production, and availability. Of 1,437
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scats collected, 996 contained remains of vertebrate prey. Most cats (80-85%) were house-based
and obtained from 15 to 90% of their food from natural prey, depending on abundance and
availability of the latter. Wild rabbits (Oryctolagus cuniculus) were the most important prey, and
cat responded functionally to changes in abundance in availability of this prey. Prolonged snow
cover made rabbits vulnerable to cats irrespective of abundance. Small rodents were the second
most important cat prey, while brown hares (Lepus europeus) and birds were less important. In a
period with high rabbit abundance, cat predation corresponded to 4% of annual production of
rabbits and to about 20% of annual production of field voles (Microtus agrestis) and wood mice
(Apodemus silvaticus). Prey choice of feral cats was similar to that of the house-based cats, but
as the former subsisted almost completely on natural prey, their absolute intake (294 g/day
during years with high rabbit abundance) was four times that of an average house-based cat (66
g/day).

Palomares, Francisco and Miguel Delibes. 1994. A note on the movements of a free-ranging
male domestic cat in southwestern Spain. Hystrix 5 (1-2): 119-123.
(Online: http://www.italian-journal-of-mammalogy.it/include/getdoc.php?
id=538&article=104&mode=pdf )

Rochlitz, I. BVSc, MSc, PhD, MRCVS. 2003. Study of factors that may predispose domestic
cats to road traffic accidents: part 1 The Veterinary Record 153(18): 549-553
Abstract: Between March 2000 and February 2001, six veterinary practices in
Cambridgeshire collected data on 117 owned cats which they had examined after a road accident.
For one week every month during the same year, the practices distributed questionnaires to the
owners of cats which had been examined for reasons other than a road accident, and the data
from these cats were checked to ensure that they were representative of the practice records,
which were compared with a survey of the owned cat population for age and sex. From this
population, the cats that were allowed outdoors and had never been in a road accident were
chosen as controls, and compared with the cats that had been in a road accident for age, sex,
pedigree status and coat colour. The cats that had been in a road accident differed from the
control population with respect to age, sex and pedigree status; for every one-year increase in
age, the odds of a road accident decreased by 16 per cent; the odds for males (entire and
neutered) being in a road accident were 1·9 times the odds for females (entire and neutered), and
the odds for pedigree cats were 0·29 those for non-pedigree cats.
Part 2. The Veterinary Record 153(19): 585-588
Abstract: Between March 2000 and February 2001, six veterinary practices in Cambridgeshire
collected data on 117 owned cats which they had examined after a road traffic accident (RTA).
The owners of 66 of these cats completed questionnaires and the information in them was
compared with the information in questionnaires from a control population of 796 cats that had
never been in a road accident. The RTA cats were, on average, younger than the controls; when
adjusted for age the two populations did not differ with respect to the time they spent outdoors or
the time they had lived at their current address. Proportionately more of the RTA cats wore
reflective collars and/or lived in areas with higher levels of traffic than the controls. The
accidents were evenly distributed throughout the year, and there was a trend for more accidents to
happen during the night than the day. Forty-eight per cent of the accidents whose location was
known occurred just outside or very near the cat's home.

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Schmidt, Paige, Roel Lopez, and Bret Collier. 2007. Survival, fecundity, and movements of
free-roaming cats. Journal of Wildlife Management 71(3):915-919
Abstract: Free-roaming cats (e.g., owned, semi-feral, and feral) impact wildlife worldwide
through predation, competition, and disease transmission. Baseline ecological information
necessary for population management is lacking. We radiocollared free-roaming cats (feral, n =
30; semi-feral, n = 14; owned, n = 10) in Caldwell, Texas, USA between October 2004 and
November 2005 and compared population demographics among sex and ownership
classification. We found ranges and movements declined across ownership classes whereas
survival and fecundity increased. Our findings suggest that human interactions (e.g., feeding)
may result in high, localized free-roaming cat densities, which may concentrate feral cat impacts
and should be considered when evaluating population control strategies.

Steen-Ash, Sara. 2004. Intraspecific spatial dynamics of urban stray cats. Shaw et. Al.
Eds: Proceedings 4th International Urban Wildlife Symposium: 222-227.
(Online: http://ag.arizona.edu/pubs/adjunct/snr0704/snr07042s.pdf )
Abstract. Supporters of the TTVAR (trap-test-vaccinate-alter-release) approach for the
control of stray cat populations assume that a group of cats sharing a common food source will
defend their space and resources from immigrating individuals. Previous studies have indicated
that group- living domestic cats exhibit a high degree of home range overlap, associate
frequently and amicably with group members, and defend their resources against immigrating
individuals. This study was designed to examine the spatial relationships among cats managed
with the TTVAR method on the Texas A&M University campus. Specific objectives of this
study include quantifying individual home ranges and examining the spatial overlap and degree
of association between individuals. Nineteen cats from 6 sites were fitted with radiocollars in
September and October 1998, and 11 cats from 5 sites were fitted with radiocollars in January
and February 1999. Males ranged an average area of 15.2 ha. The mean female home range size
was 12.8 ha. No significant difference was found in home range size between males and females
(P< 0.3244, Mann-Whitney U). In all 6 sites, cats exhibited a high degree of home range overlap;
however, only 15 individuals were found to associate with other cats. The majority of associating
pairs of individuals were found together infrequently. The findings of this study suggest that
most cats living on the Texas A&M University campus do not exhibit the same spatial dynamics
as expected from colonies of individuals sharing a common food source. Behaviors common to
cats living in cohesive groups were observed in this population of cats only occasionally.
Consequently, the assumption of resource defense by individuals sharing a common feeding area
may not fully apply for this population of stray cats.

Tennent, Jaclyn and Colleen T. Downs. 2008. Abundance and home ranges of feral cats in
an urban conservancy where there is supplemental feeding: a case study from South
Africa. African Zoology 43(2):218-229.
Abstract: There is much debate surrounding the impact of feral cats (Felis catus) on wildlife.
Conservancies are usually areas where indigenous flora and fauna are protected and aliens
excluded or managed. The University of KwaZulu-Natal's Howard College campus (HCC) is an
urban conservancy containing feral cats that are presently not managed, and little is known about
their ecology and behaviour. Consequently a feral cat population census was conducted, and
their home range investigated. Estimates of the overall campus feral cat population numbers
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ranged between 23.4–40.0 cats/km2 with a minimum of 55 identified as resident. They were not
randomly distributed in the study area, with spacing patterns being related to resource
availability. Home range area and core distribution of eight radio-collared cats were determined
over 13 months. Total home range areas were relatively small, with considerable overlap
between them. Home ranges were clustered in areas with permanent feeding stations and these
were also within the cats' core ranges. Supplemental food resources appear to have a major
influence on numbers, home and core range area, and behavior of cats. It is clear that cat
densities grow to high levels with reliable and abundant food supply and only ad hoc
sterilization. This has implications for their management in the HCC urban conservancy.

Turner, D. and Mertens C. 1986. Home range size, overlap and exploitation behavior in
domestic farm cats (Felis catus). Behaviour 99: 22-45.
Abstract. Home range size, overlap and exploitation in domestic farm cats is examined. Data
were collected on the 11 adult cats (5 ♂♂; 6 ♀♀) living on 3 Swiss farms throughout 1984; 8 of
the cats were fitted with radio transmitters. Census-point and focal animal methods of data
collection were employed. From area-observation curves we concluded that our home range
maps (and sizes) were good approximations of the areas utilized by the animals in 1984.
Correlations existed between range size and 1) the number of map-fields visited per day by the
cat (+); 2) the percentage of observations at the primary home (-); 3) the proportion of the range
visited each day (-); and 4) the percentage of map-fields used only by one cat (+, but only for
males). The general pattern of social organization found elsewhere was confirmed: males were
generally more tolerant of each other than females (based on range overlap), especially
considering animals living on different farms. Animals from the same primary home showed
considerable range overlap. Male ranges were much smaller than expected (not even twice as
large as the female ranges, whereas they should have been about 10 times the size, based on all
other studies). These results are discussed in connection with male reproductive strategies, cat
density and patterns of range utilization. And lastly, we could demonstrate coordination of
hunting activity in time and space by two adult individuals on one farm (sibs), adding to the list
of behavior patterns illustrating facultative sociality of house cats.

Warner, Richard. 1985. Demography and movements of free-ranging domestic cats in

rural Illinois. Journal of Wildlife Management. 49:340-346
Abstract: Free-ranging domestic cats on farmsteads were censused annually in August 1977-
81 within a 5,182-ha area typical of the cash-grain region in central Illinois. The estimated
average number of cats on the area in late summer was 326 (6.3/100 ha). Annual recruitment of
immature cats into the late summer population averaged 1.5/adult female. Survival beyond 3-5
years of age was rare; <1% survived 7 or more years. Eleven adult cats were radio-monitored
during a 30-day period in summer; four males ranged over larger areas (P < 0.01) than seven
females (228 ± 100 ha and 112 ± 21 ha, respectively). When cats were not on farmsteads,
approximately 73% of their radiolocation points (N = 1,227) were in edge or linear
configurations of cover. Cats made disproportionately high (P < 0.05) use of farmsteads and
perimeters, roadsides, and field interfaces and disproportionately low use (P < 0.05) of fields of
corn and soybeans. Domestic cats on the area were well fed by humans but routinely deposited
prey at their residences.

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BOOKS

Gehrt, Stanley, D., Seth P.D. Riley, and Brian L. Cypher, editors. 2010. Urban Carnivores:
Ecology, Conflict, and Conservation. The John Hopkins University Press.

Johnson-Bennett, Pam. 2004. Cat vs Cat: keeping the peace when you have more than one
cat. New York, NY: Penguin Books.

Leyhausen, Paul. 1979. Cat Behavior: the predatory and social behavior of domestic and wild
cats. Garland STPM Press.

Tabor, Roger. 2003. Understanding Cat Behavior. Cincinnati, OH: F&W Publications Inc.

Turner, Dennis. C., and Patrick. Bateson, editors. 2000. The Domestic Cat: the biology of its
behavior. New York, NY: Cambridge University Press.

WEBSITES

Cats International
http://www.catsinternational.org/index.html
Article on territories:
http://www.catsinternational.org/articles/natural_cat_behavior/cats_view_of_territory.html

California Fish and Game

http://www.dfg.ca.gov/wildlife/nongame/nuis_exo/dom_cat/dfgcatbiblio.html
A bibliography of feral, stray, and free-roaming domestic cats in relation to wildlife
conservation. 1994. Compiled by Ronald M. Jurek. Calif. Dep. of Fish and Game, Nongame
Bird and Mammal Program Rep. 94-5. 24 pp.

Center for Companion Animal Health

http://www.vetmed.ucdavis.edu/CCAB/strays.htm
Bibliographies, websites and resources on free-roaming cats and dogs.

Know Your Cat

Article on outdoor cat territories: http://www.knowyourcat.info/info/teritory.htm

Recommended articles and websites

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