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In many countries, high densities of domestic cats (Felis catus) are found in urban habitats where they have the potential to exert considerable predation pressure on their prey. However, little is known of the ranging behaviour of cats in the UK. Twenty cats in suburban Reading, UK, were fitted with GPS trackers to quantify movement patterns. Cats were monitored during the summer and winter for an average of 6.8 24 h periods per season. Mean daily area ranged (95 % MCP) was 1.94 ha. Including all fixes, mean maximum area ranged was 6.88 ha. These are broadly comparable to those observed in urban areas in other countries. Daily area ranged was not affected by the cat's sex or the season, but was significantly larger at night than during the day. There was no relationship between area ranged and habitat availability. Taking available habitat into account, cat ranging area contained significantly more garden and other green space than urban habitats. If cats were shown to be negatively affecting prey populations, one mitigation option for consideration in housing developments proposed near important wildlife sites would be to incorporate a 'buffer zone' in which cat ownership was not permitted. Absolute maximum daily area ranged by a cat in this study was 33.78 ha. This would correspond to an exclusory limit of approximately 300–400 m to minimise the negative effects of cat predation, but this may need to be larger if cat ranging behaviour is negatively affected by population density.
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Ranging characteristics of the domestic cat (Felis catus)
in an urban environment
Rebecca L. Thomas &Philip J. Baker &
Mark D. E. Fellowes
#The Author(s) 2014. This article is published with open access at Springerlink.com
Abstract In many countries, high densities of domestic cats (Felis catus) are found in urban
habitats where they have the potential to exert considerable predation pressure on their prey.
However, little is known of the ranging behaviour of cats in the UK. Twenty cats in suburban
Reading, UK, were fitted with GPS trackers to quantify movement patterns. Cats were
monitored during the summer and winter for an average of 6.8 24 h periods per season. Mean
daily area ranged (95 % MCP) was 1.94 ha. Including all fixes, mean maximum area ranged
was 6.88 ha. These are broadly comparable to those observed in urban areas in other countries.
Daily area ranged was not affected by the cats sex or the season, but was significantly larger at
night than during the day. There was no relationship between area ranged and habitat
availability. Taking available habitat into account, cat ranging area contained significantly
more garden and other green space than urban habitats. If cats were shown to be negatively
affecting prey populations, one mitigation option for consideration in housing developments
proposed near important wildlife sites would be to incorporate a buffer zonein which cat
ownership was not permitted. Absolute maximum daily area ranged by a cat in this study was
33.78 ha. This would correspond to an exclusory limit of approximately 300400 m to
minimise the negative effects of cat predation, but this may need to be larger if cat ranging
behaviour is negatively affected by population density.
Keywords Domestic cat .Felis catus .Buffer zone .Ranging area .GPS
Introduction
The UKs pet cat (Felis catus) population is in excess of 10 million individuals (Murray et al.
2010) with an additional 800,000 feral cats (Harris et al. 1995), thereby far exceeding
populations of any other mammalian carnivore (Harris et al. 1995). As pet cats receive
supplemental food from their human owners, their densities do not reflect that of their prey,
but are instead influenced by housing density (Sims et al. 2008;Thomasetal.2012)and
socioeconomic status (Murray et al. 2010). Therefore, cat populations may be exceptionally
Urban Ecosyst
DOI 10.1007/s11252-014-0360-5
R. L. Thomas (*):P. J. Baker :M. D. E. Fellowes
School of Biological Sciences, Harborne Building, University of Reading, Whiteknights, Reading,
Berkshire RG6 6AS, UK
e-mail: rebecca.thomas@reading.ac.uk
high in some urban areas (2001,500 cats km
2
:Libergetal.2000; Baker et al. 2008; Sims
et al. 2008;Thomasetal.2012).
Domestic cats are opportunistic predators and will readily depredate what is available
(Turner and Bateson 2000), responding to prey density and availability (Fitzgerald and
Turner 2000). As a result of this behaviour and their artificially maintained high densities,
domestic cats have the potential to exert considerable predation pressure on their prey (Loss
et al. 2013). In one study of prey returned home by free-ranging urban domestic cats, 11
mammal, 21 bird and two reptile species were returned, with one species, the wood mouse
(Apodemus sylvaticus) accounting for 40 % of all prey (Thomas et al. 2012). Although the
numbers and diversity of prey taken by pet cats have been widely investigated (Churcher and
Lawton 1987;Barratt1997b,1998;Robertson1998; Woods et al. 2003; Baker et al. 2005,
2008; van Heezik et al. 2010;Thomasetal.2012;Loydetal.2013) the degree to which they
may be negatively affecting wildlife populations in urban settings is still equivocal (Crooks
and Soulé 1999; Lilith et al. 2010). Furthermore, while most studies have considered the direct
effects of cat predation, other studies have emphasised the potential negative indirect conse-
quences of domestic cats (Beckerman et al. 2007; Bonnington et al. 2013), further depressing
prey numbers in areas with high cat densities.
Areas important for biodiversity conservation in many developed countries are increasingly
being encroached upon by urban development (e.g. for the UK, Goddard et al. 2009)andthe
issue of cat predation has been raised as a potential problem in housing developments
constructed near these areas. Some developers have discussed a ban on cat ownership on
new developments built in close proximity to important conservation areas, but many believe
that such bans would not be enforced (Leake and Cracknell 2006). Understanding the
movement and behaviour of domestic cats is important in understanding their potential
predatory effect (s) within an area, especially as a cats range may encompass areas of high
wildlife value. For example, Lilith et al. (2008) suggested a buffer zone of 360 m around any
housing development in Australia to reduce the numbers of pet cats hunting in important
wildlife areas. Such figures may, however, not be directly translated to other countries because
of inter-national differences in urban landscape structure and, perhaps, differences in the
abundance and types of prey species available. Additional data are, therefore, required on
the range size of urban cats in different countries.
An animals home range will usually contain areas in which it will be able to find food, rest
and reproduce (Burt 1943). Free-ranging pet cats are, however, rather unusual in that they are
fed and provided with shelter at the same point such that their ownershomeislikelytobea
strong focal element in their ranging behaviour. The cat may also be neutered, so they may not
be driven by a strong desire to mate. Furthermore, range size may be negatively affected by cat
density and the dispersion of natural food sources (Liberg et al. 2000).
Most research into home range and territory use of animals has been performed using radio
tracking (Attuquayefio et al. 1986;Konecny1987;Meek2003;Harper2007;Hucketal.2008;
Morgan et al. 2009). Although this technique has been widely used, there are well known
issues with accuracy and the degree of resolution (Schmutz and White 1990), particularly in
some habitats. Furthermore, tracking also frequently involves a human researcher directly
following a focal individual such that it is possible that the act of tracking itself might affect the
animals behaviour (Maclean 2007). This may be particularly problematic with companion
animals which might be expected to exhibit both attraction and avoidance responses to
humans. One method for circumventing these problems, and gaining higher resolution spatial
data, is through the use of global positioning systems (GPS).
Current estimates of home range sizes have focused mainly on feral or semi-feral cats
(Liberg 1980;Apps1986; Konecny 1987;Pageetal.1992; Mirmovitch 1995; Barratt 1997a;
Urban Ecosyst
Edwards et al. 2001; Naidenko and Hupe 2002;Biroetal.2004;Harper2007; Schmidt et al.
2007; Recio et al. 2010) which are likely to have larger ranges than pet cats as they have to
provision themselves (Barratt 1997a). Consequently, these will contain multiple food resources
which may change in abundance throughout the year (Liberg et al. 2000;Harper2007)
possibly resulting in changes in range size, position or utilisation (Schmidt et al. 2007). In
contrast, those of pet cats will be focused on their owners home to which they return regularly
to feed and rest throughout the day and night. As urban cats can attain higher densities than
feral cats it is important to understand their movement behaviour and whether habitat
complexity affects this trait (Horn et al. 2011).
Previous studies of the movements of urban domestic cats have been carried out in the
USA, Australia and New Zealand, but with no previous studies in the UK (Table 1). Collec-
tively, these have indicated no significant differences in range size between males and females
(Barratt 1997a;Meek2003;Morganetal.2009; Metsers et al. 2010; van Heezik et al. 2010)or
between summer and winter seasons (Morgan et al. 2009), but an increase at night (Barratt
1997ba; Metsers et al. 2010: but see van Heezik et al. 2010). The complexity of habitat use by
cats and the effect of habitat availability on ranging behaviour has not been investigated
previously. Therefore in this study we used GPS tracking to obtain estimates of the area ranged
by free-ranging pet cats in a large English town and how these varied in relation to (1) season,
(2) sex, (3) day versus night and (4) habitat availability.
Methods
Study area and participants
The study was conducted in Reading, Berkshire, UK (51°27N, 0°58W) during summer (6th
July-6th August 2009) and winter (18th January-18th February 2010) seasons. Reading is a
large town, 40 km west of London, which covers an area of c.55km
2
and has a population of
approximately 230,000 people. Twenty cats (15,5) from 19 households were recruited by
leaflet dropping and door-knocking (Thomas et al. 2012). All cats in the study were neutered
and had unrestricted access to the outdoors through a cat flap.
Data collection
GPS receivers (CatTrack, Perthold Engineering, USA) were used to record cat movement
behaviour. Units were attached using a specially designed harness, so that the receiver was
positioned on the back of each cat, behind the shoulder blades. Each harness was fitted with a
safety break-away clip so that it would come off if the cat became entangled. GPS units
weighed 22 g (4.4×2.7×1.3 cm) and the harness approx. 4045 g, < 5 % of each cats body
mass, and were powered by a lithium battery.
GPS receivers were programmed to record a location every minute, although when unable
to get a signal they would not record until a signal was re-established. All GPS systems will
have some form of error when calculating location points. The error for this system was
calculated in a number of different conditions (indoors, outside under cover, outside clear)
along a predetermined 1 km route in June 2009. The deviation of each point from the route was
measured in ArcGIS Version 9.3 (ESRI 2011) and on average recorded within an accuracy of
6 m (range 010 m), although when indoors many fixes were missed.
Each cat was tracked for up to eight 24 h periods in each season, with a minimum of 24 h
between tracking periods to minimise ownersconcerns about the wearing of the harness for
Urban Ecosyst
Tab le 1 Mean daily area ranged (DAR, 95 % minimum convex polygon [MCP]) and mean maximum daily ranged (MDAR, 100 % MCP) presented for present study and other
studies of urban cat movement patterns
Source Location Length
of study
Method
1
N
2
Mean (minimummaximum)
daily area ranged
(ha;95 %MCP)
Mean (minimummaximum)
maximum daily area ranged
(ha;100 %MCP)
Results of statistical
comparisons
between
3
:
Seasons Sexes Day/night
Current study Reading, UK 2 months GPS 20 1.94(0.994.23) 6.88(1.6911.15) NS NS S (DAR) NS
(MDAR)
Barratt (1997a) Canberra, Australia 9 months RT 10 5.31(0.0227.93) 8.58(0.0243.56) NS S
Kays and DeWan (2004) Albany, USA 4 months RT 11 0.24(0.011.30) 0.65(0.133.0) ––
Meek (2003) Jervis Bay, Australia 16 nights RT 14 2.92(0.0414.65) NS
Morgan et al. (2009) Christchurch, New Zealand 12 months RT 21 1.8(0.110) NS NS
Metsers et al. (2010) Canterbury, New Zealand 10 days GPS 38 26(372) NS S
van Heezik et al. (2010) Dunedin, New Zealand 6 days GPS 32 3.2(0.4821.75) NS NS
1
Method: GPS global positioning system; RT radio tracking
2
Number of individual cats studied
2
NS not significant; Ssignificant; denotes not tested
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long periods of time. Cats were monitored by their owners during deployment and devices
were removed immediately if any signs of distress were observed, although this did not happen
during the course of the study.
Location data were downloaded from the device and were converted for use in ArcGIS
Version 9.3 (ESRI 2011). Location points from the first 30 min were removed, as during this
period the owner would have been fitting and re-adjusting the harness.
Data analysis
Ranging area
Ranging areas were estimated using both 95 % and 100 % minimum convex polygons (MCPs)
using Home Range Extension software (Rodgers et al. 2007). MCPs were used, first, to allow
comparisons with previous studies and, second, because of the temporally correlated nature of
these data. 95 % MCPs were used as a measure of the area typically covered during each 24 h
period (hereafter daily area rangedor DAR) and 100 % MCPs were used as a measure of the
maximum daily area ranged (MDAR). Each location was classified as day or night, calculated
using the daily sunrise and sunset times. MCPs were calculated for each day sampled and the
mean was used to get an average 95 % and 100 % MCP for each cat within each season.
Land use data were derived from the Ordnance Survey Mastermap collection (EDINA,
University of Edinburgh). Each daily 95 and 100 % MCP was cut from the Mastermap layer,
and the resulting areas for each habitat category were extracted for each range. Twenty one
habitat categories occurred within the study site, and these were combined to form three broad
categories: urban land use (buildings, roads etc.), green land use (grassland, woodland, etc.)
and private gardens (Appendix 1).
To normalise data for analysis, mean MCP areas were square root transformed. All data
analyses were undertaken using Genstat (12th edition). Day and night effects and seasonal
effects on range area were analysed using paired sample t-tests, and sex effects were analysed
with unpaired t-tests.
Habitat effects
The relationship between habitat availability (measured as habitat available within a 350 m
radius of the cats home; distance based on figures from Lilith et al. (2008) and results of this
study) and the habitat composition of the cats DAR was analysed in two ways. First, Jacobs
D Index (Jacobs 1974) was used. Jacobs Index is calculated as D = (r-p)/(r + p 2rp),where
D provides an indication of how habitat availability influences preference, r is the proportion
of habitat utilised and p is the proportion of habitat available. D ranges from +1 (complete
preference), through 0 (no influence) to 1 (complete avoidance). The 95 % confidence
intervals are calculated for D; if 0 is not included in the range, then a significant (P<0.05)
habitat preference or avoidance is supported.
Second, compositional analysis was performed following Aebischer et al. (1993) and
Dickson and Beier (2002). Here, additive log ratios were calculated for garden and green
habitats, the proportion of urban habitat as the denominator. These ratios were calculated for
both habitat available and habitat used; if there is no habitat preference then the difference of
the ratios will not be significantly different from zero. An overall comparison is made by
calculating Wilkslambda (Λ), then transforming this into the test statistic (=N×ln(Λ)) where
N is the number of cats studied. Degrees of freedom are calculated as the number of habitat
types available 1. The test statistic approximates the χ
2
distribution. If a significant (p<0.05)
Urban Ecosyst
overall effect is found, then relative habitat use can be compared using a series of paired t-tests
(Dickson and Beier 2002).
To test whether range size was influenced by habitat availability (within 350 m radius of
home) or habitats used (in DAR), Pearson correlation analyses were performed between
ranging characteristics (day and night DAR and MDAR respectively) and the difference of
the log ratios of habitats used.
Results
Twenty cats were tracked in the summer session, with 14 (11,3) continuing through
to the winter. The loss of cats was due to cat deaths, householders moving home, or
householders withdrawing from the study. The average age of the study cats was
7.4 years (range 116 years). Of the eight 24 h tracking periods each season, the mean
number completed by each cat was 7.0 (range: 38) and 6.7 (range: 38) in summer and
winter, respectively; on average, those cats tracked in both seasons completed 11.7
(range: 1116) tracking periods (Table 2).
Tab le 2 Mean daily (95 % minimum convex polygon [MCP]) and maximum daily (100 % MCP) area (ha)
ranged by each cat tracked during the course of the study. Columns headed maximum denote largest range area
recorded for each individual
Cat No. of days
tracked
Daily area ranged (95%MCP)(ha)Maximum daily area ranged (100%MCP)(ha)
Mean [95%CI]Maximum Mean [95%CI]Maximum
1 6 4.23 [2.765.71] 6.24 8.22 [4.8211.62] 16.28
215 1.95[1.412.49] 4.71 8.83 [6.1811.48] 17.54
315 1.10[0.821.37] 2.07 4.14 [2.365.92] 11.69
413 1.27[0.871.66] 2.44 3.37 [2.154.59] 7.80
511 2.13[0.793.32] 3.32 9.10 [3.6314.57] 33.78
610 2.26[1.962.56] 3.32 7.61 [4.6910.52] 16.67
714 2.66[1.733.58] 7.55 8.34 [5.4311.26] 23.03
8 5 0.99 [0.731.25] 1.34 2.74 [1.164.31] 5.65
9 8 1.51 [1.281.74] 2.07 8.78 [6.3411.21] 15.55
10 8 1.0 [0.741.26] 1.52 1.69 [1.212.16] 2.61
11 14 2.22 [1.602.84] 5.49 8.33 [5.6810.97] 21.67
12 16 1.87 [1.532.21] 3.24 10.59 [7.7513.43] 26.89
13 5 1.68 [1.192.17] 2.61 5.45 [2.957.96] 8.43
14 16 2.77 [2.393.16] 4.62 10.58 [8.3212.83] 22.70
15 12 1.23 [0.971.50] 1.99 3.20 [2.324.08] 5.89
16 3 1.23 [1.031.43] 1.43 3.53 [3.383.68] 3.67
17 5 3.68 [2.754.60] 5.42 11.15 [8.9313.37] 13.80
18 12 1.70 [1.232.16] 3.36 4.16 [2.296.04] 10.11
19 16 1.87 [1.692.06] 2.55 5.37 [4.136.61] 11.02
20 12 1.92 [1.642.19] 2.98 7.07 [5.748.40] 10.77
Mean 10.8 1.94 [1.802.09] 3.41 6.88 [6.217.55] 14.28
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Ranging area and maximum ranging area
No significant differences were found in the daily area ranged (paired t-test: 95 % MCP;
t
13
=1.64, p= 0.124) or the maximum daily area ranged (100 % MCP; t
13
=1.23, p=0.241)
between seasons. As no seasonal differences were evident, the data have been combined in
subsequent analyses.
No significant differences were found between males and females with respect to daily
(independent sample t-test: t
19
= 1.36, p= 0.190) or maximum daily (t
19
=1.31,p= 0.207) area
ranged. Mean DAR was 1.94 ha (95 % CI: 1.802.09 ha); mean MDAR was 6.88 ha (95 % CI:
6.217.55 ha) (Table 2). Maximum values for DAR and MDAR were 7.55 ha (mean: 3.41 ha)
and 33.78 ha (mean: 14.28 ha) respectively (Table 2).
There was a significant difference in the mean daily area ranged during the day (1.84 ha)
and night (2.74 ha) (paired t-test: t
19
=4.66, p< 0.001). The result for maximum daily area
ranged approached significance (t
19
=2.03, p= 0.057).
Habitat effects
Analysis of habitat selection showed that cats had more garden and green habitats in their
ranging area that expected; urban habitats were avoided (p< 0.05 for each; Fig. 1). Formal
compositional analysis supported this result (χ
2
= 28.16, d.f. = 2, p< 0.001). Detailed inspec-
tion showed that there was a significant preference for garden (paired t-test: t
19
= 5.23,
p< 0.001) and green habitats over urban (paired t-test: t
19
=2.78,p= 0.012), but no significant
difference between relative amount of green and garden habitats in ranging areas, when
availability was considered (paired t-test: t
19
=0.48, p= 0.64).
As range areas differed significantly between day and night, their relationships with habitat
availability and use were investigated separately. As there was no significant difference in
relative garden and green space use, these were considered together and the log-ratio of both to
urban habitat was used as the explanatory variable. There was no significant effect of habitat
used or habitat available on DAR or MDAR (all p>0.2).
0
0.1
0.2
0.3
0.4
0.5
0.6
Garden Green Urban
Mean (+/-se) habitat available / used
Habitat
Fig. 1 Mean (+/SE) proportion of habitat available within 350 m radius of owners house (stippled bars) and
proportion of habitat in ranging area (open bars) of study cats
Urban Ecosyst
Discussion
In this study, ranging areas were broadly comparable to those from other countries
(Table 1). Similarly, we did not detect any seasonal differences (Morgan et al. 2009),
even though cats might be expected to range further in summer due to more favourable
weather conditions, nor between the sexes (Barratt 1997a;Meek2003;Morganetal.
2009; Metsers et al. 2010; van Heezik et al. 2010). While it should be noted that the
sample size was small, with a low number of female cats, they lie within the typical
range of sample sizes for such studies (Barratt 1997a; Meek 2003;KaysandDeWan
2004). Furthermore, the cats utilized in this study were all neutered. In the UK this is
the norm, with over 91 % of adult domestic cats sterilised (Murray et al. 2009;Thomas
et al. 2012).
Significant differences in ranging areas between day and night were found and near
significant differences were also found in day and night maximum ranging area. Cats
ranged further during the night (see also Barratt 1997a; Metsers et al. 2010) but other
studies have found no difference (van Heezik et al. 2010). Although domestic cats
show a greater tendency towards diurnal activity than feral cats, possibly due to their
domestication (Turner and Bateson 2000), feral cats are more active at night (Alterio
and Moller 1997). This study shows that pet cats still may exhibit this tendency, by
ranging further. This may be in part due to decreased road traffic and less conspecific
conflict, as evidence has shown the spatial movement patterns of cats are determined
by busy roads and density, spatial distribution and social dominance of individual cats
(Barratt 1997a).
There were no significant relationships between the daily area ranged, maximum daily area
ranged and habitat type. This is unexpected, as an a priori expectation may be that cats would
roam over a wider area if less suitable habitat was available. However, cats did vary in habitat
preferences, spending disproportionately more time in gardens and other green habitats, than
urban habitats. Taken together, these results suggest that while habitat availability does not
affect the ranging area, it does influence habitat use, with cats spending more time in gardens
and green areas. This in itself is unsurprising, given that ranging cats will be seeking prey and
avoiding disturbance.
It is vital when considering the effects of cat predation on wild bird and small
mammal populations to understand how cats use their environment. When discussing
the mitigation techniques that could be implemented, the banning of cat ownership
near ecologically sensitive areas, such as heathland, has been proposed (Leake and
Cracknell 2006). The acceptability of this measure to residents within the UK is
mixed though, with less than half of respondents questioned feeling that this is a
justifiable approach (Thomas et al. 2012). This may become more acceptable to the
public once it is determined whether or not cats are detrimentally affecting wildlife
populations.
Lilith et al. (2008) has suggested a buffer zone concept, whereby developments, or
households owning cats, may be banned within a certain distance from an area of
important wildlife value. Data from the radio-tracking of rural and urban cats in Australia
produced a figure of 360 m that would be acceptable as a buffer zone (Lilith et al. 2008).
Within our study, the largest maximum daily area ranged by a cat was 33.78 ha, this
being equivalent to a circular area of diameter of 656 m. However, ownershomes are
not generally located at the extreme periphery of their pets ranges, such that the radius
(328m)maybeamoreusefulmetricforestimatingtheappropriatesizeforabuffer
zone. Furthermore, it could be argued that is an overly conservative approach. For
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example, estimates based on the mean MDAR (6.88 ha) and mean DAR (1.94 ha) would
be substantially smaller, radii of 148 m and 79 m respectively, and caution should also be
taken due to the small sample size in this study.
One additional factor that may also affect the ranging behaviour of cats that has not been
considered in this study is the density of other cats and whether domestic cats are quasi-
territorial (feral cats typically are semi-social and territorial: Genovesi et al. 1995;Halletal.
2000). If it were the case that range size was negatively affected by cat density, then higher
densities of cats may decrease range sizes and indeed this may explain the differences in
nocturnal maximum ranging size considered above. It would be important to understand these
effects, as it could be the case that by reducing numbers of cats in urban areas it may increase
the areas ranged by those that remain; such density-dependent responses would suggest that
buffer zones would need to be larger than those identified from studies of existing cat
populations. Again this would suggest that managers should err on the side of caution.
Domestic cats are the most common mammalian predator in the UK, and as a result of their
close links with people, cat densities can be exceptionally high in urban areas. With the recent
growth in interest in urban ecology, it has become clear that many populations of birds and
mammals are on the edge of sustainability in these human-modified habitats (e.g. Blair 2004;
Goddard et al. 2009), as urban animals respond to the challenges of living in highly altered and
fragmented habitats. While native predators are still present in these habitats (albeit in lower
densities, e.g. sparrowhawk Accipiter nisus; Chamberlain et al. 2009), human behaviour results
in exceptionally high populations of an introduced predator, the domestic cat. This study, the
first of its kind in the UK, demonstrates that to be effective in safeguarding important wildlife
habitats, any zone of restricted cat ownership would need to be in the order of 300400 m.
Acknowledgments We wish to thank the cat owners and cats who took part in this research. RLTwas supported
by a NERC doctoral training award. Requests for original data may be made to the corresponding author.
Open Access This article is distributed under the terms of the Creative Commons Attribution License which
permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are
credited.
Appendix 1
Original land-use categories and details of their re-classification.
Urban land use Green land use Private Gardens
Manmade surface or step Natural surface Gardens
Path Coniferous trees
Road Coniferous-scattered
Road traffic calming Nonconiferous trees
Structure Nonconiferous-scattered
Track Orchard
Unclassified Rough grassland
Unknown surface Scrub
Archway Inland water
Building
Greenhouses
Urban Ecosyst
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Urban Ecosyst
... Our estimates of home ranges for non-boundary cats are similar to previous estimates from the large nearby conurbation of Reading Thomas, Baker, & Fellowes, 2014). In contrast, the mean home range (95% KDE) of our boundary study cats was ~ 70% larger than that of our non-boundary cats (3.42 ha vs 2.01 ha). ...
... m; individual maximum 1502 m) cats, suggesting that cats were equally capable of movement, but that their likelihood of movement differed. The influence of sex, with males having larger ranges, supports some studies (Kays et al., 2020) but not others (Thomas et al., 2014). While cat age was present in the home range model it was not significant, which mirrors Hanmer, but c.f. Hall et al., 2016). ...
... A buffer zone > 300 m around natural areas (328 m, Thomas et al., 2014;360 m, Lilith, Calver, & Garkaklis, 2008), outdoor cat enclosures (catios) and cat curfews have been suggested as management options, with the latter implemented by some Australian councils (Council, 2019). Although catios appear to be gaining popularity and confining cats or restricting access would help to reduce predation while benefiting cat welfare (Grayson & Calver, 2004), their introduction in the UK would be met with strong opposition due to the restricting of roaming behaviour being perceived as cruel (McDonald, Maclean, Evans, & Hodgson, 2015). ...
Article
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The domestic cat (Felis catus) is a predator of global significance. In Great Britain there are ca. 9.5 million owned pet cats, with their population determined by human population density. As urban areas expand and encroach on areas of conservation value, it is not known how cats use these areas and how habitat availability influences predation rates. To address this, over a year we recorded the movement and prey of 79 owned cats in inner suburban areas (non-boundary cats) and in areas adjacent to natural habitats on the edge of the suburban area (boundary cats). Boundary cats had larger home ranges (mean 3.42 S.E. ± 0.61 ha) and returned more prey (mean 7.91 S.E. ± 2.70 prey cat⁻¹year⁻¹) than cats in non-boundary areas (2.01 S.E. ± 0.70 ha; 3.35 S.E. ± 1.06 prey cat⁻¹year⁻¹respectively). Assuming a prey return rate of 23%, extrapolated predation rates equate to 34.40 (S.E. ± 11.74) and 14.57 (S.E. ± 4.62) prey cat⁻¹year⁻¹ in our boundary and suburban study sites respectively. While non-boundary cats had little access to natural habitats, natural habitats made up > 25% of the home range of boundary cats. Boundary cats travelled a mean distance of 64.9 m (S.E. ± 6.8) into these natural habitats, with some cats ranging > 300 m inside these areas. Bird predation rates did not differ between boundary and non-boundary cats, but boundary cats killed three times more mammals. This is of relevance to urban planning, as the hunting behaviour of pet cats extends the ecological effects of urbanisation into surrounding habitats.
... In the United States, United Kingdom, Australia, New Zealand, and other countries, there is an ongoing debate about whether cats should be allowed to freely roam outdoors [2]. Concerns about the ecological impact of cats on wildlife populations and native species, feline transmission of zoonotic diseases, and the inconvenience that roaming cats might cause to other people have led to various suggested initiatives, including night curfews, collars with bells, cat-free buffer zones around nature reserves or sensitive conservation areas, and routine confinement [3][4][5][6][7][8][9][10]. Another factor supporting indoor confinement is the concern that free-ranging cats may be injured by other cats, humans, cars or predators (e.g., feral dogs, coyotes, wolves) [11]. ...
... The most common method used to understand the movement patterns of domestic cats is the estimation of home ranges. Home ranges have been estimated in a number of countries beyond Denmark, including Sweden [27], Norway [33,34], France [24], the United Kingdom [2,8,10,35], Switzerland [36], Corvo Island [31], the United States [2,30,[37][38][39], Africa [40], Australia [2,4,7,17,41], and New Zealand [2,5,6,22,26,29,42,43]. ...
... In France, suburban cats have been found to have a larger home range during April to June [24]. On the other hand, temporal variation was not confirmed in other studies [8,9,25,30,31]. Cats may enlarge their home ranges in spring in response to changes in the weather and available prey. It is possible that seasonal variation was already represented in the dataset we analysed, since the lowest temperature in the tracking period was −5.2 • C and the highest temperature was 26.9 • C (mean temperature = 11.9 • C). ...
Article
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We studied the roaming patterns of companion cats in Denmark. The movements of 97 cats with outdoor access were traced for about seven days using GPS tracking. Data on the cats were gathered from their owners. The median time cats spent away from their homes was 5 h per day (IQR: 2.5 to 8.8 h), median daily distance moved was 2.4 km (IQR: 1.3 to 3.7 km), and median for 95% BBKDE home range was 5 ha (IQR: 2.9 to 8.5 ha). Cats above seven years of age spent less time away from home, were less active and had a smaller home range than younger cats. Cats with access to nature areas spent more time away from home, were more active and had larger home ranges. Intact male cats spent more time away from home than neutered cats and had larger home ranges as well. Finally, rainfall had an impact on the distance moved by cats: on days without rainfall the cats moved 3.6 km on average (95% CI: 2.8; 4.5 km); and on days with heavy rainfall the cats moved 2.4 km on average (95% CI: 1.6; 3.5 km).
... Partial curfews tend to be more acceptable to owners, with nocturnal mammals being the main beneficiaries of nighttime confinement of cats (Woods et al., 2003), while night-time or crepuscular confinement, particularly in warmer months, is recommended when most wild species are active and in their reproductive periods (Mori et al., 2019). Some studies have shown that cats with unrestricted outdoor access roam significantly further at night than during the day (Metsers, Seddon, & van Heezik, 2010;Thomas, Baker, & Fellowes, 2014), while others found no differences (Hanmer et al., 2017;van Heezik et al., 2010). Similarly, whether a cat was kept indoors at night or allowed unregulated access to the outdoors has previously been found to have no impact on home range size (Hall, Bryant, Fontaine, & Calver, 2016). ...
... Cats that were allowed unrestricted outdoor access had 75% larger home ranges, reached 46% greater maximum distances from home, and showed 31% greater daily total distances traveled than cats with partially restricted outdoor access. The greater extents of ranging displayed by the cats that were free to roam by day and night, likely stem from significant differences between day and night in cat ranging areas (Thomas et al., 2014). The number of prey brought home was not related to the extent of home ranges, suggesting that cats tend to roam, at least in part, to fulfill behaviors unrelated to hunting. ...
... In terms of the spatial extent of domestic cat ranging, home ranges and core ranges were small and broadly consistent with those of domestic cats in other studies (Castañeda et al., 2019;Hall, Bryant, Haskard, et al., 2016;Kays et al., 2020). In contrast to other studies, which have found that older cats had smaller home ranges than younger ones (Castañeda et al., 2019;Hall, Bryant, Haskard, et al., 2016;Kays et al., 2020), and males tended to have bigger home ranges than females (Kays & DeWan, 2004;Thomas et al., 2014), our model did not find significant effects of these variables on home ranges, though older cats did tend to stay closer to home and travel less far on a daily basis. Cats living in households in rural or urban settlements, with a wide range of human population densities, showed similar spatial behavior in this study. ...
Article
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Domestic cats (Felis catus) that roam outdoors have increased exposure to hazards to their health and welfare. Outdoor cats can themselves present a hazard to biodiversity conservation and wild animal welfare. Approaches to reducing predation of wildlife by cats might also bring benefits to cats by reducing their roaming and associated risks. We investigated ranging behaviors of domestic cats that regularly captured wild prey, and that had restricted or unrestricted outdoor access. We tested whether interventions aimed at reducing predation also affected their spatial behavior. We evaluated cat bells, Birdsbesafe collar covers, using a “puzzle feeder”, provision of meat‐rich food, object play, and a control group. Seventy‐two cats in 48 households in England completed the 12‐week trial in spring 2019. Home ranges were small (median AKDE95 = 1.51 ha). Cats with unrestricted outdoor access had 75% larger home ranges, 31% greater daily distances traveled, and reached 46% greater maximum distances from home, than cats with restricted outdoor access. None of the treatments intended to reduce predation affected cat ranges or distances traveled. While owners might use interventions to reduce predation, the only effective means of reducing cat roaming and associated exposure to outdoor hazards was restriction of outdoor access. While some interventions, like object play and dietary changes, can reduce domestic cat predation of wildlife, we found that these measures do not affect cat roaming behavior. Restricting access to the outdoors, even partially, does reduce the extent of cat ranging and likely reduces associated exposure to outdoor hazards.
... Десятки научных публикаций посвящены численности, распределению и хищничеству кошек в населенных пунктах (Barratt, 1998;Koenig et al., 2002;Мерзликин, 2003;Lepczyk et al., 2003;Baker et al., 2010;van Heezik et al., 2010;Balogh et al., 2011;Gehrt et al., 2013;Loss et al., 2013;Loyd et al., 2013;Shipley et al., 2013;Bartos Smith et al., 2016;Loss, Marra, 2017;Malpass et al., 2018 и др.). Особенно озабочены этими вопросами британские исследователи (Fitzgerald, 1988;Woods et al., 2003;Baker et al., 2008;Beckerman et al., 2007;Sims et al. 2008;Thomas et al., 2012Thomas et al., , 2014Bonnington et al., 2013;Aegerter et al. 2017 и др.). Действительно, в английских городах много домашних, имеющих хозяев, кошек, причем их плотность варьирует более чем на порядок: значения свыше 200 особей/км 2 -обычное дело, а максимальные могут превышать 2500 особей/км 2 (Baker et al., 2008;Sims et al., 2008;Thomas et al., 2012;Aegerter et al. 2017). ...
... Кроме того, само присутствие кошек оказывает сублетальные и опосредованные летальные воздействия на репродуктивный успех некоторых видов птиц (Beckerman et al., 2007;Bonnington et al., 2013). По некоторым сведениям, в урболандшафтах городские домашние, имеющие хозяев, кошки предпочитают проводить больше времени на озеленных участках с неплотной застройкой, нежели в собственно городских "биотопах" (например, Thomas et al., 2014). Однако вместе с тем, они избегают внутренних частей крупных озелененных и природных территорий (см. ...
... Owner reports (N = 210) about the reason why their cats no longer wear a collar also included collar incidents such as scratching and skin irritation (4%), entrapment of jaw, mouth, or tongue (2%), and entrapment on objects (2%) ( Lord et al., 2010 ). In a survey among British cat owners (N = 113), 26% of owners stated that they had removed a collar because of injury or signs of distress ( Thomas et al., 2014 ). ...
Article
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There are multiple reasons for cats to wear collars, among the most relevant are identification and mounting of radiofrequency tracking devices or predation deterrents. Reports on severe incidents with cat collars indicate that both, entrapment of the collar on an object or a body part, have the potential to lead to serious injury or death. Therefore, expert opinions and guidelines concerning the safety of cat collars are controversial. This review focuses on the current state of knowledge weighing welfare risks against potential benefits of collar use in domestic cats (Felis catus). The results of this review show that even “safety collars” (collars with a breakaway clip or elastic parts) do not fully prevent severe incidents, albeit rarely reported. Nevertheless, the use of breakaway collars is considered vital. To further reduce risk of entrapment, good fit (i.e., the collar must not be too loose nor too tight), undamaged material and close monitoring during adjustment periods are recommended. Behaviors such as excessive scratching or rubbing and coat or skin problems such as matting, alopecia or erythema were reported in association with collars. These adverse effects on behavior, coat or skin can be manageable in pet cats. The propensity of a breakaway collar to open/release varies between types of collars and brands. For unsupervised use, a collar that opens/releases easily is recommended to reduce risk of entrapment. However, this feature can limit the benefits of collar use as the intended functions of the collar might no longer be maintained. The main benefits provided by a collar are visible identification and mounting of radiofrequency tracking devices (to locate cats via GPS) or predation deterrents (to reduce the impact of predation on wildlife). Given the risks associated with collars, there are no benefits to cats wearing collars solely for “fashion”. Comparatively, collars have a lower inherent risk than other scenarios cats encounter while free-roaming. Yet risks and benefits regarding collar use should be assessed for each individual case. Future directions of research should focus on ways to further reduce or prevent risks of collars. One interesting research direction is whether training cats to tolerate collars can reduce discomfort and risk of entrapment or skin irritation. Paper collars have not been investigated yet, but are cheap and can be replaced at low cost, they can be inscribed with owner contact details and tear easily.
... Indeed, individual cats may also compete with one another as density increases, thus affecting per capita predation rates Barratt 1998; Thomas et al. 2012; but see Lepczyk et al. 2004). However, in many countries, high densities of pet cats are found in urban habitats, resulting in a considerable predation pressure on their prey (Thomas et al. 2014;Legge et al. 2020). Additionally, even if the predation mortality were null, small reductions in fecundity or growth rate due to fear could result in a substantial decrease in bird abundance, especially in urban areas where cat densities are high (Beckerman et al. 2007;Grade et al. 2021). ...
Article
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The domestic cat (Felis catus) is one of the most abundant predators and a serious threat to many wildlife species. While a large body of literature explores the number and diversity of individuals depredated by pet cats, the drivers of predation have been investigated much less. Although the environment of the cat, the owner behavior, and the intrinsic characteristics of the cat itself could impact the predatory behavior and should therefore not be considered separately, very few studies simultaneously take these three components into account. In this study, we explored 21 concomitant drivers of predation by pet cats linked to these three components at different scales, to explain the owners-reported frequencies of captured birds, mammals, and herpetofauna. Among the 1,400 sociological surveys received from cat owners, 740 reliable answers were analyzed. Results suggest that the owners-reported prey capture frequencies were strongly influenced by the environment, especially by factors relating to urbanization. Rural owners were around two times more likely to report more frequent predation events than owners living in urban areas, whatever the group of prey studied. As a result, the urban habitat variable had the highest impact on predation in this study. An experimental approach would be beneficial to identify the factors influencing the reported predation rates, which are causally related to the number of wild animals killed.
... One way to gain insight into how animals use urban spaces is through habitat selection analyses to assess habitat preferences and / or avoidances within the context of landscape-scale distribution or home range utilisation (Saunders et al. 1997;Dowding et al. 2010a; Thomas et al. 2014;Roberts et al. 2017;Mueller et al. 2018). However, understanding habitat use on a finer scale can yield greater benefits in conservation planning (Gilioli et al. 2018), particularly for species that perceive the environment at small spatial scales (Ritchie and Olff 1999), have limited dispersal ability (Gilioli et al. 2018) and / or which may be associated with specific habitats or microhabitats (Banks and Skilleter 2007). ...
Article
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Understanding patterns of habitat selection and factors affecting space use is fundamental in animal conservation. In urban landscapes, such knowledge can be used to advise householders on how best to manage their gardens for wildlife. In this study, we tracked 28 West European hedgehogs ( Erinaceus europaeus ), a species of conservation concern in the UK, in an area of high-density housing using radio and GPS tags to quantify patterns of habitat use and identify factors associated with the proportion of time spent in individual gardens. Both males and females exhibited a preference for residential gardens, but there were subtle differences between the sexes in relation to house type and front versus back gardens. Hedgehogs spent significantly more time in gardens where artificial food was provided, where a compost heap was present, if foxes ( Vulpes vulpes ) were infrequent visitors, if it rained overnight and as daylength increased (i.e., shorter nights); garden use was not significantly associated with variables potentially likely to reflect invertebrate prey abundance. These data suggest that the primary positive action that householders can undertake for urban hedgehogs is providing supplementary food. However, householders often feed hedgehogs after they know they are already visiting their garden. Consequently, the presence of artificial food may make it difficult to identify other important influences affecting garden use. Finally, we report that a GPS fix acquisition rate < 60% likely had no major effect on the results of our analyses, but should be a consideration in future studies using this technique on this species and in this habitat.
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a) The Landscape of Fear (LOF) concept proposes that a prey individual’s perceived risk of predation can affect their distribution and activity. Prey may perceive the risk of predation following the detection of predator cues, such as auditory, visual, and chemical signals; this has been demonstrated in numerous studies. b) There is limited understanding on the extent of the role that chemical predator cues play under the LOF concept. Within this study, a range of olfactory cues, including those of native and invasive predators, were applied to artificial feeders used by eastern grey squirrels (Sciurus carolinensis) in a single urban site in the United Kingdom. Motion- activated cameras recorded squirrel feeding and vigilance behaviours. The durations of such behaviours were collated and analysed to determine whether the simulated presence of predators using affected squirrel foraging. c) Domestic cat (Felis catus) odours significantly reduced proportional squirrel feeding duration at feeders, compared to passive and active controls (distilled water and rabbit (Oryctolagus cuniculus domesticus) urine, respectively). Pine marten (Martes martes) faeces significantly reduced proportional squirrel feeding duration compared to feeding under passive control applications.(Werner and Anholt 1993; Maynard-Smith et al. 2003; Scott-Phillips 2008) d) Our results confirm previous knowledge on the LOF theory; olfactory cues of a predator disrupt prey foraging. These observations provide an insight into how predator scents can affect the distribution, feeding, and anti-predation behaviours of prey. e) Our results are discussed in the context of the following areas: the application of predator scents as a deterrent of domestic cats in an urban environment; the wider effects of pine marten reintroductions on introduced eastern grey squirrel populations; the evolutionary significance of predator chemical cues.
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We took advantage of cat regulations enacted within differing subdivisions in the City of Armadale, Western Australia, to test the hypotheses that the species diversity (measured by the Shannon-Weiner index) and abundance of small and medium-sized mammals should be higher in native bushland within or adjacent to subdivisions where cats are restricted compared to similar areas where cats are not restricted. There were three different regimes of cat regulation: no-cat zone (strict prohibition of cat ownership applying in one site), compulsory belling of cats and night curfew at one site, and unregulated zones (free-roaming cats applying at two sites). Both sets of cat regulations were in place for approximately 10 years prior to our survey. We also measured structural and floristic features of the vegetation at each site that might influence the species diversity and abundance of small and medium-sized mammals independently or interactively with cat activity. No significant differences in species diversity were found across the sites and KTBA (known-to-be-alive) statistics for Brushtail Possums Trichosurus vulpecula and Southern Brown Bandicoots lsoodon obesulus, the two most abundant medium-sized mammals present, were similar across all sites. The smaller Mardo Antechinus flavipes, which could be regarded as the most susceptible to cat predation of all the native species trapped because of its size, was trapped mostly at an unregulated cat site. Total mammals trapped at the unregulated cat sites exceeded those caught at the two sites with restrictions, but these unregulated sites also had significantly denser vegetation and there was a borderline (p = 0.05) rank correlation between vegetation density and mammal captures across all sites. It appears that pet cats are not the major influence on the species diversity or abundance of small and medium-sized mammals at these sites and that vegetation characteristics may be more important.
Article
The 14 contributions, 9 abstracted separately, which explore the behaviour and ecology of Felis catus (= Felis silvestris catus) are arranged in 4 major sections (development of young cats, social life, predatory behaviour, cats and people), together with an introduction and a postscript. -P.J.Jarvis
Article
Seven males and one female of house cat (Felis silvestris var. catus) were radiotracked in Solling, 50 km to the north-west of Göttingen, Germany, during 1995 and 1997-1998. 4955 locations (2417 in 1995 and 2538 in 1997-1998) were used for analysis. The home range size of the house tomcats was 90.2-294 hectares (MCP-method for 100% of radio-lacations without obvious deviations) or 20.3-175.1 hectares (95% adaptive Kernel analysis). Seasonal changes in the home range use were analyzed under estimation of overlapping the territories during time intervals. During a year, the 1 : 4 to 1 : 51 rations were found as minimum and maximum sizes of the monthly home ranges in the tomcats. Significant seasonal changes in the use of home range by tomcats were described, including the mating season and the changes caused by human activities. The home range use of males was the most flexible during the mating season and the most stable in October-December. The mean daily activity of males was 44.5%, maximum is 58.2% in the mating season. The cats were more active and used larger area during nighttime than in daytime. The home ranges of neighboring tomcats overlapped each other; sometime, home range of one tomcat covered the whole home range of other male. Howerer, the cats used their home ranges independently of the conspecifics, except for the mating period. The distribution of food and shelters determined the habitat use by the tomcats. The tomcats did not use forest areas (the main habitat of European wild cat in Solling).
Article
Using gut samples, faecal analysis, records of prey brought home by house cats and uneaten remains in the field, the diet of domestic and feral Felis catus is examined. In descending order of frequency, mammals, birds and (especially below latitude 35o) reptiles predominate. Cat predation on islands, where bird prey is proportionally more significant, often has an adverse impact on native species. Diet is discussed in terms of sex and age differences; seasonal variations; and prey availability. The impacts of cats on farmyard rats; on wild rabbits Oryctolagus cuniculus, voles and other rodents; on game species; on bird populations on continents; and on island wildlife, are all discussed. -S.J.Yates
Chapter
Discusses patterns in density and home range size, there being great variability in each for both sexes in Felis catus. Group living is usually associated with females and kittens, though adult males are sometimes included. Roaming behaviour and overlap in male home ranges are noted. Mating tactics are discussed: there is no active mate choice in females, who mate with the most dominant male present; and although close kin matings are not uncommon, inbreeding is often avoided by females in oestrus temporarily leaving groups which contained related males. Comparison is made with spatial behaviour in other felids. -S.J.Yates
Article
We used Monte Carlo simulations to investigate the effects of animal movement on error of estimated animal locations derived from radio-telemetry triangulation of sequentially obtained bearings. Simulated movements of 0-534 m resulted in up to 10-fold increases in average location error but <10% decreases in location precision when observer-to-animal distances were <1,000 m. Location error and precision were minimally affected by censorship of poor locations with Chi-square goodness-of-fit tests. Location error caused by animal movement can only be eliminated by taking simultaneous bearings.