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The Impact of Anthropogenic Factors on the Behavior, Reproduction, Management and Welfare of Urban, Free-Roaming Cat Populations


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Free-roaming domestic cats constitute an integral part of many urban ecosystems worldwide; their presence results from undisturbed natural reproduction, abandonment by pet owners, and abundant food resources. These cats present controversial and emotional issues regarding management of their high densities, hygiene and epidemiologic risks to humans, and predation of wildlife. Improved urban cat management requires greater knowledge of the anthropogenic factors affecting free-roaming cat populations. In this study, we explored the relationship between caretaker treatment levels and the socio-economics of the caretakers and neighborhoods, and free-roaming cat reproduction management, behavior, and physiology in urban populations. Eight free-roaming cat feeding groups, from eight neighborhoods in Tel Aviv, Israel, were observed for six months. Neighborhoods displayed differences in socio-economic status and housing type, and cat feeding groups reflected different levels of caretaker treatment with regard to neutering, physical interaction with the cats, and medical treatment. We examined whether agonistic behaviors, neutering rates, pregnancy rates, and cortisol levels of the cats differed between groups. We found a strong socio-economic effect on neutering and pregnancy rates: low-income neighborhoods had lower neutering and higher pregnancy rates. Higher-level caretaking was associated with lowered aggression, as well as lower cortisol levels of neutered females. Additionally, analysis of data from 622 registered cat feeding groups from north (high-income) and south (low-income) Tel Aviv, obtained from the municipal veterinary department, revealed a socio-economic influence on reproduction management. Our results indicate that, in urban environments, both the neighborhood's socio-economic status and the caretaker treatment level affect management of these cat populations. We conclude that, to improve cat management success, municipalities should consider addressing these socio-economic differences, while, just as importantly, raising awareness and encouraging caretaker involvement in neutering efforts. Improved caretaker treatment levels can lead to reduced cat aggression, with consequent improved cat welfare and reduced noise disturbance to humans.
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The Impact of Anthropogenic
Factors on the Behavior,
Reproduction, Management
and Welfare of Urban, Free-
Roaming Cat Populations
Hilit Finkler
, Erez Hatna
and Joseph Terkel
Zoology Department, George S. Wise Faculty of Life Sciences, Tel
Aviv University, Tel Aviv 69978, Israel
Institute for Management Research, Radboud University, The
ABSTRACT Free-roaming domestic cats constitute an integral part of many
urban ecosystems worldwide; their presence results from undisturbed natural
reproduction, abandonment by pet owners, and abundant food resources.
These cats present controversial and emotional issues regarding management
of their high densities, hygiene and epidemiologic risks to humans, and preda-
tion of wildlife. Improved urban cat management requires greater knowledge of
the anthropogenic factors affecting free-roaming cat populations. In this study,
we explored the relationship between caretaker treatment levels and the socio-
economics of the caretakers and neighborhoods, and free-roaming cat repro-
duction management, behavior, and physiology in urban populations. Eight
free-roaming cat feeding groups, from eight neighborhoods in Tel Aviv, Israel,
were observed for six months. Neighborhoods displayed differences in socio-
economic status and housing type, and cat feeding groups reflected different
levels of caretaker treatment with regard to neutering, physical interaction with
the cats, and medical treatment. We examined whether agonistic behaviors,
neutering rates, pregnancy rates, and cortisol levels of the cats differed between
groups. We found a strong socio-economic effect on neutering and pregnancy
rates: low-income neighborhoods had lower neutering and higher pregnancy
rates. Higher-level caretaking was associated with lowered aggression, as well
as lower cortisol levels of neutered females. Additionally, analysis of data from
622 registered cat feeding groups from north (high-income) and south (low-
income) Tel Aviv, obtained from the municipal veterinary department, revealed
a socio-economic influence on reproduction management. Our results indicate
that, in urban environments, both the neighborhood’s socio-economic status
and the caretaker treatment level affect management of these cat populations.
31 Anthrozoös DOI: 10.2752/175303711X12923300467320
Address for correspondence:
Hilit Finkler,
Zoology Department,
George S. Wise Faculty of
Life Sciences,
Tel Aviv University,
Ramat Aviv, Tel Aviv 69978,
We conclude that, to improve cat management success, municipalities should consider addressing
these socio-economic differences, while, just as importantly, raising awareness and encouraging care-
taker involvement in neutering efforts. Improved caretaker treatment levels can lead to reduced cat
aggression, with consequent improved cat welfare and reduced noise disturbance to humans.
Keywords: caretakers, free-roaming cats, population management, socio-economic status
(SES), TNR, urban ecosystem
Free-roaming domestic cats constitute an integral part of many urban ecosystems
worldwide. Their presence is the result of many years of undisturbed natural repro-
duction, abandonment by pet owners, and an abundance of food resources, inten-
tionally or unintentionally supplied by the human population (Strula 1993; Natoli 1994; Gunther
and Terkel 2002). The management of these cats is a controversial and emotional issue, lead-
ing to concerns about cat welfare (Natoli 1994), hygiene and epidemiologic risks to the human
population (Slater 2001), potential predation risk to wildlife (Clarke and Pacin 2002; Jessup
2004), and the best methods of control (Wallace and Levy 2006).
The co-existence of free-roaming cats with humans in the urban ecosystem has been stud-
ied from numerous perspectives, such as cat activity patterns (Haspel and Calhoon 1993), food
preferences and availability (Church, Allen and Bradshaw 1996; Bradshaw et al. 1999), and cat
and feeder interrelationships (Natoli et al. 1999; Centonze and Levy 2002). However, a much
smaller number of studies (Calhoon and Haspel 1989; Alves et al. 2005; Schmidt, Lopez and
Collier 2007) has incorporated anthropogenic factors, attempting to solve issues related to free-
roaming cat overpopulation. Such incorporation of human factors was recently acknowledged
by several authors as crucial to the efficient management of human–wildlife conflicts in urban-
ized environments (Baker and Harris 2007; Markovchick-Nicholls et al. 2008), and in the case
of managing the cat population, it is assumed to play an equally important role.
Part of the free-roaming cat population in urban areas is cared for, to varying extents, by vol-
untary cat caretakers: from food delivery alone, to comprehensive care including neutering and
medical treatment (Haspel and Calhoon 1990; Centonze and Levy 2002). The caretakers de-
liver food to feeding groups, where the cats gather to feed and where they interact (Izawa 1983;
Natoli, De Vito and Pontier 2000). The extent of contact between these caretakers and cats,
and the selective behaviors that caretakers choose to engage in, can vary, with probable con-
sequent varying effects on the cat population (Toukhsati, Bennett and Coleman 2007). Some
caretaker behaviors are likely to contribute to the maintenance of the free-roaming cat popula-
tion, as, for example, many of them engage in feeding but not in neutering (Natoli et al. 1999;
Centonze and Levy 2002; Toukhsati, Bennett and Coleman 2007). In addition to the effect of
caretaker behaviors, the particular socio-economic status (SES) of a neighborhood or region of
a city can also affect the cat population. For example, the socio-economic status of neighbor-
hoods surrounding urban parks in Phoenix, Arizona, was recently shown to predict bird species
richness and abundance in those parks (Kinzig et al. 2005; Shaw, Chamberlain and Evans
2008). Furthermore, Beck (2002) showed that in Baltimore, Maryland, more free-roaming dogs
were found in poor neighborhoods compared with wealthier ones, partially due to the former
having more available refuse and open trash cans in the streets.
If cat behavior, reproduction, and welfare parameters are affected by the behaviors of hu-
mans (the caretakers of a feeding group), the measures of such parameters in the cats should,
to some extent, consequently reflect the cultural, social, or economic differences pertaining to
their caretakers. In addition, larger-scale effects on free-roaming cats, as on other animals,
32 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
Finkler et al.
derive from city-level management decisions (Kinzig et al. 2005). In order to better understand
the effects of anthropogenic factors on these cat feeding groups, we therefore selected sev-
eral parameters of cat reproduction, behavior, and physiology, and measured them against the
different types of neighborhood housing, the socio-economic status of the caretakers and
neighborhood, and the level of caretaking.
Our four hypotheses were as follows:
1. Frequency of agonistic behaviors of the cats before and during food delivery would be
influenced by behaviors of the caretakers, and also, to a smaller extent, by the neigh-
borhood housing type, which reflects density of human presence and consequent level
of human activity.
2. Neutering rates would be largely influenced by city-level management of the cat popu-
lation, but might also be influenced by the involvement level of the caretaker him/herself.
3. Pregnancy rates would be affected by the same factors affecting neutering rates.
4. Cortisol levels, as a stress indicator, would be affected by the level of caretaking as
well as by the SES of the neighborhood, which could affect the general attitude of res-
idents towards the cats.
Thus, this research is the first step in achieving a greater understanding of the impact of
anthropogenic factors on free-roaming, urban cat populations and their relevance to cat pop-
ulation management.
Tel Aviv is the main city of the greater Tel Aviv metropolitan area and Israel’s largest urban re-
gion, ca. 53 km² (Tel Aviv population census of 1995–1998). The study areas were located in
high SES residential neighborhoods of north Tel Aviv, and low SES residential neighborhoods
of south Tel Aviv (Porat, Frenkel and Shoshany 2008) (northern neighborhoods in the survey
comprised ca. 12 km² and southern neighborhoods ca. 15 km²). The process of selecting
low- and high-SES neighborhoods for the study is elaborated below. In stage 1 of the study,
we examined whether these two contrasting SES areas differed in cat feeding group disposi-
tion and characteristics. In stage 2, we compared reproductive, behavioral, and physiological
parameters of cats from feeding groups located in eight different neighborhoods, four in north
and four in south Tel Aviv.
Stage 1: Characteristics of Cat Feeding Groups in Tel Aviv
Active cat feeding is a widespread phenomenon throughout the city. The municipal veterinary
services cooperate with the cat caretakers in neutering the cat populations, and 40,000 cats are
reported to have been neutered since 1994 (Z. Galin DVM, Tel Aviv chief municipal veterinarian,
personal communication). Following cat caretakers’ requests, the animals are taken by the vet-
erinary services from the feeding group for a TNR (Trap-Neuter-Return) procedure. Each cat en-
tering the procedure is registered according to the feeding group from which it was taken, and its
sex, age, sexual status, and any injuries are noted. The tip of the left ear is cut, as in other coun-
tries (Natoli et al. 2006), to allow identification of neutered cats. After neutering, cats are returned
to their group. This procedure is done at no cost to the caretaker. The relevant data from the Tel
Aviv Veterinary Services for 2000–2005 were uploaded to an Excel database and were subse-
quently used to compare the following parameters between north and south Tel Aviv: (1) number
33 Anthrozoös
Figure 1. Scores for each of the Tel Aviv neighborhoods from the factor analy-
sis. Neighborhood numbers are neighborhood identifiers from the population
census of 1995–1998. The eight neighborhoods selected for the study are
marked by arrows (on the lower right side, southern low-SES neighborhoods,
and on the upper left side, northern high-SES neighborhoods).
of feeding groups, (2) number of neutered cats, (3) number of visits made to feeding groups by
the veterinary municipal services, (4) rates of pregnancies, and (5) rates of rabies vaccinations.
Stage 2: Field Study
Selection of Neighborhoods: According to the most recent Tel Aviv population census (1995–
1998), Tel Aviv comprises 63 neighborhoods. To select the neighborhoods for this study, five
socio-economic variables were determined from the population census for each neighbor-
hood (matriculation rates, employment rates, number of new immigrants, computer ownership,
and income level) and entered into a factor analysis (using the principal axis factoring extrac-
tion method in SPSS). This analysis takes all the variables and produces, according to the
correlation between them, a smaller set of factors. The first factor in the analysis explained
75% of the variance in the data, and was interpreted as the socio-economic status. All five vari-
ables had a loading of 0.74 or above on the first factor. The other factors had eigenvalues
lower than 1 and were therefore not considered further. The scores of the neighborhoods from
the first factor were used to choose the neighborhoods for the study.
Of the 63 neighborhoods, 49 were analyzed (there were missing data for the other 14). Fig-
ure 1 shows the neighborhood scores, produced by the factor analysis on the SES data (the first
factor in the factor analysis), for the Tel Aviv neighborhoods. Neighborhoods that received a low
score were labeled “low SES” and those that received a high score were labeled “high SES.” Fi-
nally, eight neighborhoods were selected according to the factor analysis (presented in Figure 2),
as well as according to the type of housing characteristic of the neighborhood (neighborhoods
with a mean of more than two stories per building were labeled “apartment-building neighbor-
hoods;” those with a mean of two or fewer stories per building were labeled “private-home neigh-
borhoods”). Table 1 summarizes the SES, the type of housing of the neighborhoods, and the five
socio-economic variables, according to which neighborhoods were assigned to low or high SES.
34 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
Figure 2. The eight selected neighborhoods and the distribution of 622
registered cat feeding groups (for 2000–2005) (dots) in north and south Tel
Aviv. The numbers are the neighborhood identifiers: 1–4 in north Tel Aviv
and 5–8 in south Tel Aviv. Map prepared using ArcView GIS 9 software.
Finkler et al.
Selection of Feeding Groups: In order to select feeding groups, cat caretakers from the eight
neighborhoods were contacted and asked if they would be willing to participate in a study con-
sisting of observing their cat groups for six months. An initial screening process consisting of a
short telephone interview with 55 caretakers was followed by 25 visits to potential feeding groups.
During the visits, the following criteria for inclusion of groups in the study were examined:
1. Adequate amount and type of food delivered to the cats (diet based on dry commer-
cial food).
2. A cooperative caretaker.
3. A group comprising at least 10 cats.
4. No objection by neighbors to our presence or to the feeding of the cats.
5. Access to all parts of the feeding site area, in order to easily observe all the cats.
35 Anthrozoös
Table 1. Socio-economic status, type of housing of the neighborhoods selected for the study, and the five socio-economic variables used in the neigh-
borhood selection process.
Neighborhood Neighborhood SES Type Income Scale* Employed New Matriculated Owns
Identifier of Housing (%) Immigrants (%) a Computer
(%) (%)
Neot Afeka Bet 1 H A 164 60.0 7.9 51.4 48.6
Afeka 2 H P 186 52.4 10.9 51.9 44.5
Ramat Aviv Gimel 3 H A 185 55.7 9.8 51.0 48.6
Neve Avivim 4 H A 169 53.0 14.9 53.0 43.0
Yad Eliyahu 5 L A 74 46.3 15.3 41.9 25.7
Hatikva 6 L P 53 44.6 26.0 35.7 17.2
Kiryat Shalom 7 L A 60 45.6 23.0 33.5 20.4
Dakar 8 L A 56 43.8 26.0 31.7 19.6
For SES: L = low, H = high.
For type of housing: P = private, A = apartment building.
Neighborhood identifiers are the same as those in Figure 2, detailing the geographic locations.
* Average monthly income per capita, normalized so that 100 is the Tel Aviv average.
36 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
Figure 3. One of the eight feeding groups in the study, showing the cats
gathering to eat from a communal food source.
Finkler et al.
According to these criteria, eight feeding groups were selected, one in each of the eight
neighborhoods of the field study. Before the start of observations, the Tel Aviv veterinary mu-
nicipal services were informed of the field study, in order to ensure continuous and well-
arranged observation periods, and to prevent any unexpected changes in neutering status of
the cats. Feeding groups were chosen with no preliminary knowledge of the proportion of
neutered cats. A typical feeding station is shown in Figure 3.
Determination of Caretaker Treatment Level: Cat caretakers from the eight feeding groups
were interviewed to determine their level of treatment of the cats. They answered five ques-
tions on a five-point Likert scale (where 1 equals the strongest negative reply and 5 equals the
strongest positive reply). The interview comprised the following questions:
1. What is your level of closeness to the cats (as expressed by individually identifying and
being able to pet or otherwise handle most of the group cats) (1 = not close at all, 5 =
very close)?
2. How often do you contact the veterinary municipal services for TNR (1 = once a year,
5 = every 3 months)?
3. How often do you pay privately to neuter your group cats (1 = never, 5 = every 3
4. How long do you stay with the cats when you deliver them food (1 = little beyond time
needed for food delivery, 5 = long beyond time needed for food delivery)?
5. Do you provide medical treatment to sick or injured group cats (1 = never, 5 = I try to
give medical assistance to every sick or injured cat)?
According to the total sum, each caretaker was classified on the caretaking scale as giving
maximal (18–25), medium (10–17), or minimal (0–9) treatment.
37 Anthrozoös
Table 2. Numbers of neutered and intact adult males and females in the eight feeding
groups in Tel Aviv during the initial period of group identification.
Feeding Intact Intact Neutered Neutered Total
Group Females Males Females Males Cats
11 114521
23 45517
37 616635
43 49420
59 83222
66 42113
7 5 16 1 1 23
87 6181142
Studied Parameters: 1) Agonistic Behaviors: During observation sessions, agonistic behaviors
were recorded using the all-occurrences recording method (Altman 1974). Each agonistic be-
havior performed by a cat during an interaction with another cat (whether this cat was on the
initiating or the receiving end of the interaction), was recorded on paper. Since presence of the
cats in the feeding groups varied among individuals, the frequencies of agonistic behaviors
were regressed as a function of the relative presence of each cat during the six months of ob-
servations. Standardized residual values from this regression were used for the statistical analy-
sis. The agonistic behaviors recorded were: slapping, chasing, vocalizations (hiss, yowl and
growl), threat displays (approach and stare), and physical attacks.
2) Pregnancy Rate: The number of pregnancies in each feeding group was recorded during
the six months of observations. The number of kittens could not be recorded accurately since
not all kittens appeared at the feeding group site.
3) Neutered Cats: After the individual cat identification process, the proportion of the total
number of cats which were observed as neutered in each feeding group was calculated from
the first month of observations.
4) Hair Collection and Extraction for the Measurement of Cortisol Concentrations: Hair samples
were collected from 101 cats. Hair sample collection, method validation, cortisol extraction, and the
determination of cortisol concentrations (pg/mg hair) are fully described in Finkler and Terkel (2010).
Experimental Procedure: Prior to the beginning of the study, cats were individually identified at each
feeding group by direct observation. The identification process included photographing them and
recording specific details (Dards 1983; Natoli 1985; Calhoon and Haspel 1989): sex, estimated age,
body size, fur pattern, and other characteristics such as eye color and deformations. For the pur-
pose of data recording, each cat was given a specific identification number. Observations in all study
groups took place twice weekly, during daytime hours, upon food delivery, for 30-min sessions, over
a total period of six months, between January and August 2007. Each observation began when the
cats started to gather at the feeding site, before the feeder initiated food delivery, and ended when
only a few cats at most were still eating. Numbers of adult individuals (> 1-year-old) identified and
recorded in each feeding group during the period of group identification are shown in Table 2 (n =
193). During the study period, new cats migrated into the groups, and therefore the eventual num-
ber of cats as given in some of the results is greater than those originally presented in Table 2.
38 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
Finkler et al.
Statistical Analysis
Regression analyses were performed to evaluate the effect of the various anthropogenic fac-
tors on the number of neutered cats and cortisol levels. The effect of the anthropogenic fac-
tors on agonistic behaviors and number of pregnancies was analyzed using non-parametric
tests. Statistical analysis was performed using SPSS 16.0 software.
Characteristics of Feeding Groups in Tel Aviv
General Characteristics: Between the years 2000 and 2005, 622 cat feeding groups were
registered by the municipal veterinary services in 46 neighborhoods in north and south Tel Aviv
(as noted earlier, central Tel Aviv was not included in this study) (Figure 2). During this period,
3,771 cats (2,052 females and 1,719 males) were trapped, neutered, and returned to their
feeding group. Only 490 (13%) of the 3,771 cats had been vaccinated for rabies. This vacci-
nation is given only if the caretaker is willing to pay for it. Of the 2,052 females cats caught dur-
ing the six-year period, 305 (about 15%) were found to be pregnant. In November to January,
the highest mean number of estrous females was recorded. Pregnancies were observed
throughout the year, but peaked in February and March.
Characteristics of North vs. South Feeding Groups: Of the 622 feeding groups, 392 feeding
groups were located in north Tel Aviv and 230 in south Tel Aviv. The density of feeding groups
was 33.45 groups/km² in the north (12 km²), compared with 14.72 groups/km² in the south
(15 km²). In northern feeding groups, 2,283 cats were neutered compared with only 1,488
cats in southern groups. In northern neighborhoods, significantly more cats were vaccinated
against rabies compared with southern neighborhoods (chi square of observed [339 and 151
respectively] vs. expected frequencies [according to the ratio of north:south neutered cats,
expected frequencies were 193.34:296.65] is 15.32, p < 0.001).
Visits by the Veterinary Municipal Services: During the veterinary visits to the feeding groups, only
the number of cats taken for neutering was recorded (and not the number of all cats present), so
the actual group size could not be determined. We therefore present the mean number of cats
SD) captured per visit, which was significantly higher in south compared with north Tel Aviv
(numbers of cats: 4.92 ± 3.44 and 3.96 ± 2.88, in south and north, respectively, t-test, t = –3.70,
p < 0.01). Significantly more visits by the veterinary municipal services were made to northern
compared with southern groups (chi square of observed [number of visits in north and south: 575
and 288, respectively] vs. expected frequencies [according to the ratio of north:south feeding
groups, the expected frequencies were 545:320] is 5.03, p < 0.05), as well as per group (visits
per group in the north: 1.39 ± 0.05; in the south: 1.26 ± 0.05, t-test for square root transformed
values, t = 2.58, p < 0.05). Of the 622 feeding groups visited for the TNR procedure between 2000
and 2005, only 127 groups received a second visit. The number of second visits to the northern
feeding groups was three times that of the southern groups (95 compared with 32).
Field Study: Caretaker Treatment Level and Neighborhood Socio-Economic Status
Since each of the eight feeding groups was located in either a high- or low-SES neighborhood, we
refer to these groups correspondingly as either high or low SES feeding groups. Of the four high
SES feeding groups, three caretakers scored between 18 and 25 points (maximal treatment level)
and one caretaker scored between 10 and 17 (medium treatment level). The mean interview score
SD) for caretaker treatment level in the northern groups was 20 ± 5.29. Of the four low SES
39 Anthrozoös
Figure 4. Agonistic behaviors of cats from the eight study groups, at each
of the caretaker treatment levels. Agonistic behavior values are the
residuals from a regression of agonistic behavior frequencies on cat
presence rates. To analyze the difference between the treatment levels,
only positive values (above the x axis) were considered. The numbers in
parentheses represent numbers of cats.
40 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
feeding groups, three caretakers scored between 0 and 9 points in the interview (minimal treatment
level), and one caretaker scored between 18 and 25 points (maximal treatment level). The mean
interview score (± SD) for caretaker treatment in the southern groups was 12.25 ± 8.53.
Field Study: Effect of Anthropogenic Factors on Behavior, Reproduction
Management, and Physiology of the Cats
Agonistic Behaviors: Agonistic behavior values (which are the residuals after regression of ag-
onistic behavior frequencies on cat presence rates) were not normally distributed. Therefore,
we used non-parametric tests to examine the effect of sex, neuter status, and anthropogenic
variables (type of housing, SES of neighborhood, and caretaker treatment level), separately, on
the frequency of agonistic behaviors.
Neutered cats showed lower frequencies of agonistic behaviors compared with intact cats
(Mann-Whitney U test, Z = –2.72, p < 0.01). Caretaking level also had a significant effect on
frequency of agonistic behaviors (Kruskal-Wallis, ² = 12.99, p < 0.01, maximal (n = 161) =
–0.18 ± 0.05, medium (n = 31) = 0.12 ± 0.18, minimal (n = 74) = 0.35 ± 0.16, see Figure 4),
and a post-hoc test revealed that aggression was higher among cats receiving the minimal
treatment level compared with those receiving the maximal treatment level (p < 0.01). To fur-
ther examine the difference between the caretaking levels, we summed the positive agonistic
behavior values for each treatment level and compared them (in relation to the total popula-
tion) between levels (Figure 4). At the minimal and medium caretaking level, more cats exhib-
ited high agonistic behavior values out of the total number in that treatment level (30 of 74 and
12 of 31 cats, respectively) compared with cats at the high caretaking level (36 of 161 cats).
The contribution of the neutered and intact cats to this high aggression was equal at all care-
taker levels (Mann Whitney U test for the minimal caretaking level, Z = –0.16, p = 0.89, for the
medium level Z = –0.48, p = 0.7, and for the maximal level, Z = –0.21, p = 0.85).
Type of housing and SES had no effect on the frequency of agonistic behaviors.
Figure 5. Number of neutered and intact cats in the study groups
(groups 1–4: H = high SES, groups 5–8: L = low SES) at the beginning of
the study.
A logistic regression was performed to determine whether neighborhood SES and type of
housing are good predictors of whether a cat will be neutered or not (caretaker level was not
used in this analysis because their involvement in TNR had already been evaluated in their in-
terviews—see Methods). Socio-economic status was found to be a significant predictor of
whether a cat would be neutered (n = 193, logistic regression, B = 1.24, SE = 0.30, R
(Nagelkerke) = 0.12, p < 0.001). Neutering of cats was 3.5 times (= Exp(B)) more likely to occur
in high-SES (north Tel Aviv) groups compared with low SES-groups (south Tel Aviv). The type
of neighborhood housing was not a good predictor of neutering status in the regression model.
Cortisol Concentrations: Multiple hierarchical regression analysis was used to examine the
effects of sex, neuter status, SES, caretaker treatment level, and type of housing on cortisol
levels. In the first block, neutering and sex were entered using the enter method. In the second
block, caretaker treatment level, SES, and type of housing were entered using the forward
stepwise method. Intact cats (n = 46) had significantly higher cortisol levels compared with
Pregnancies: The frequencies of pregnancy were not normally distributed. Therefore, we ex-
amined the effect of type of housing, SES of neighborhood, and caretaker treatment level,
separately, on the frequency of pregnancies, using non-parametric tests. We found that in
high-SES groups, the frequency was significantly lower compared with the low-SES groups
(Mann-Whitney U Test, Z = –2.31, df = 64, p < 0.05). The mean number of pregnancies
SD) was 0.6 ± 0.12 (n = 23) and 0.95 ± 0.09 (n = 41) in high and low-SES feeding groups,
respectively. Type of housing and caretaker treatment level did not significantly affect
pregnancy frequencies.
Proportion of Neutered Cats: At the beginning of the study, the proportion of neutered cats of
the total number of cats in all the high SES feeding groups was higher than 50% (a range of
58–90% neutered cats). However, while three of the four low-SES groups had neutering pro-
portions of 8–23%, one low-SES group had a similar proportion to those found in the high-
SES groups (69% neutered cats) (Figure 5).
Finkler et al.
41 Anthrozoös
Figure 6. Cortisol levels in neutered females (a) intact females (b)
neutered males (c), and intact males (d) as a function of caretaker
treatment levels. No hair samples were obtained from neutered males
at the low caretaking level, and therefore are absent from the figure.
The results of this study support our hypothesis that anthropogenic factors, specifically
socio-economic status and caretaker treatment level, influence the reproduction manage-
ment, behavior, and welfare of urban free-roaming cats. Only a small number of studies
(e.g., Oppenheimer 1980; Calhoon and Haspel 1989; Childs 1990; Haspel and Calhoon
1990; Alves et al. 2005) have considered the effect of human behaviors and actions on free-
roaming, urban cat overpopulations. The present study adds to this body of knowledge
and, together with the feeding stations survey, constitutes one of the few rigorous, aca-
demic attempts to evaluate the efficacy of allocating selectively targeted financial and man-
power resources in order to achieve the most effective management of free-roaming cat
populations in Israel.
Descriptive Characteristics of Feeding Groups in Tel Aviv
The number of registered feeding stations is considerably high (622 over a six-year period) for
a city the size of Tel Aviv (53 km²). Divided by the total combined area of the northern and
neutered cats (t
= –3.47, n = 55, p < 0.001, R² = 0.14). Since the effect of caretaker
treatment level was close to significant (t
= 1.96, p = 0.053), and since the four combina-
tions of sex and neutering status showed different trends for cortisol as a function of care-
taker treatment level (see Figure 6), we tested the effect of this factor on cortisol levels in each
combination (of neutering sex) separately. Cortisol levels in neutered females with minimal
caretaker treatment level were significantly higher compared with those in neutered females
with medium and maximal caretaker treatment levels (minimal: 7.91 ± 5.22 pg/mg hair,
medium: 1.45 ± 0.38 pg/mg hair, maximal: 1.89 ± 0.79 pg/mg hair, Figure 6, one-way ANOVA,
F = 3.78, p < 0.05). Cortisol levels of intact females, intact males, and neutered males did not
differ significantly between caretaker treatment levels.
42 Anthrozoös
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
Finkler et al.
southern neighborhoods (27 km²), the mean number of feeding stations is 23/km². Natoli et
al. (2006) reported 3,533 registered colonies in Rome’s D area (517 km²) (Azienda USL Roma
D) over a period of four years, which reflects a colony density of about 7 stations/km². This dif-
ference may simply reflect the higher population density in Tel Aviv (7,500 people/km²) (Israeli
Central Bureau of Statistics 2008) compared with Rome (2,500 people/km²) (ISTAT, Instituto
Nazionale Di Statistica 2005). Beck (2002) suggests that urban stray dog population densities
are higher in areas where human density is higher, perhaps due to more abundant resources.
We suggest that where human density is higher, more food is also available to the cats, as more
people feed cats for a given area, and the result is a higher density of feeding stations in Tel
Aviv compared with Rome.
Most registered feeding groups (for TNR) were only visited once by the veterinary serv-
ices during the surveyed six-year period. Without continuous neutering, the proportion of
neutered cats in such groups will quickly become insignificant due to the immigration of in-
tact cats (Levy and Crawford 2004), and the potential long-term efficacy of this method on
a city-scale is thus jeopardized. In addition, the mean neutering rate found in our groups
was 50%. Previous authors have suggested that in order to manage a population suc-
cessfully, an annual neutering rate of at least 75% should be continuously maintained
(Xemar, Pontier and Artois 1998; Andersen, Martin and Roemer 2004). Thus, although the
city-operated TNR program is a step in the right direction, it may not be enough in itself for
efficient cat management in a city with such a successful free-roaming cat population as
Tel Aviv.
Rabies is a major zoonotic disease in Israel, and many measures are taken to prevent human
exposure to the disease (Gdalevich et al. 2000). It is therefore surprising that only 13% of the sur-
veyed cats were vaccinated and that such vaccination is not mandatory. However, this number
is probably an under-estimate, since it only represents the cats from registered colonies. As far
as optional vaccination for rabies is concerned, we are not aware of references to other studies
of free-roaming cats and therefore cannot compare our own results with other findings. Com-
pared with the optional vaccination, in other studies that initiated or followed TNR programs, ra-
bies vaccination was part of the routine procedure (Levy, Gale and Gale 2003; Wallace and Levy
2006), as the name of the program implies (TTVARM: Trap-Test-Vaccinate-Alter-Return-
Monitor) (Hughes and Slater 2002).
The proportion of pregnant females in the survey was about 15%. This figure is identical
to that found in a 12-year survey of 103,643 feral cats from seven TNR programs in the United
States (Wallace and Levy 2006). Pregnancies were mainly observed between January and
September, peaking in February and March, similar to previous reports (Nutter, Levine and
Stoskopf 2004; Wallace and Levy 2006). Lower rates of pregnancies were observed in the fall
months, between October and December, similar to other studies (Nutter, Levine and Stoskopf
2004; Wallace and Levy 2006).
A greater number of females than males was found in the current study, which was
also reported by Wallace and Levy (2006) and in a survey in England of 192 feeding
groups (Rees 1981; Remfry 1981). The reason for this skewed ratio is not clear. Perhaps
the possibility of capturing females is higher than that of capturing males, because fe-
males usually roam less than males and are therefore more likely to be present at the
feeding site (Liberg and Sandell 1988; Mirmovitch 1995). In addition, due to their greater
territorial behavior and roaming habits, male mortality may be higher than that of females
(Remfry 1981).
43 Anthrozoös
SES Effects on Reproduction and Neutering Rates and the Relationship between
the Veterinary Services and Caretakers
Several important differences were revealed in the current study between north (high SES)
and south (low SES) Tel Aviv in the characteristics, dispersion, and number of cat feeding
groups, as well as in neutering and pregnancy rates. According to our survey, feeding group
density in north Tel Aviv was more than two times that of south Tel Aviv. However, the mean
human population density does not differ between the northern and southern neighborhoods
(population census of 1995–1998 of Tel Aviv). Since the urban cat is directly dependant on hu-
mans for its food supply, whether this is supplied intentionally (i.e., by caretakers) or uninten-
tionally (i.e., from refuse or trash cans), we expect both geographical areas to be similar
“food-rich environments” (Mirmovitch 1995) and thus support a similar number of cats. In
south Tel Aviv, where fewer feeding stations are registered, the unintentional food supply may
be of greater importance for the cats. In 2006, the Green Forum of Tel Aviv, of the Society for
the Protection of Nature in Israel, issued a report on environmental health risks in south Tel Aviv.
One of the findings was that there are more large, collective trash cans and fewer small, indi-
vidual trash cans in the area compared with north Tel Aviv. The former are several meters long
and are usually open, and thus can easily accommodate the nutritional needs of several cats.
It is possible that “natural” feeding groups form around these cans (as opposed to the “artifi-
cial” ones fed by caretakers). The former are probably not registered by the TNR service, since
they do not have a daily caretaker, and thus although the number of cats in the south might
be equal to their number in the north, there are fewer registered feeding stations.
High SES is often correlated with higher education levels (Cohen, Doyle and Baum 2006), and
has also been found to be linked to higher awareness or concern regarding various health and
social issues (de Walle and de Jong-van den Berg 2007; Morley et al. 2008). Cat caretakers in
the north, due to their higher education levels, may thus have been more aware of the cat man-
agement possibilities and, consequently, more involved in neutering activities, through register-
ing their feeding group and requesting repeated veterinary visits, as well as engaging in
privately-funded TNR (resulting in more neutered cats, as was found in the field study). Several
reports suggest that the higher the education level, the higher the trust people have in the gov-
ernment or civil services (Christensen and Leaegreid 2002; Delhey and Newton 2002). South Tel
Aviv residents, of lower education levels, may have a lesser degree of trust in the TNR public serv-
ice, which is expressed in lower registration rates of feeding stations. Indeed, from in-depth in-
terviews conducted by the authors (Finkler and Terkel in press), some south Tel Aviv caretakers
reported their lack of trust as a reason for not using the veterinary public services for TNR (i.e.,
saying that the veterinary department takes the cats but does not return them), as was also
noted by caretakers in studies conducted in Rome (Natoli et al. 1999; Natoli et al. 2006).
Neutering rates were higher and reproduction rates lower in feeding groups located in high-
SES areas compared with low-SES areas. It is possible that in cases where caretakers had a
long wait for the arrival of the TNR veterinary services team, as was previously reported (Na-
toli et al. 2006), more resourceful caretakers, from high-SES neighborhoods, had neutered
the cats at their own expense, as was found in the current study. In addition to the cost of neu-
tering, another potential cost—for rabies vaccination—may pose a financial burden for the
less affluent caretaker. A single rabies vaccine is offered to the caretaker at a cost of NIS 12
(ca. US$3), and can be given to the cat during anesthesia for TNR, with the caretaker’s con-
sent. Less affluent caretakers were probably more reluctant to pay for the rabies vaccine—
hence, the lower number of vaccinations in the south.
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
44 Anthrozoös
Residents of lower SES neighborhoods may be discriminated against in terms of access
to certain municipal and social services (Williams 1998; Altschuler, Somkin and Adler 2004).
In the present study, the number of veterinary visits per cat group in northern groups was sig-
nificantly higher compared with the southern groups. It is possible that the veterinary services
responded differently to the requests received from south and north Tel Aviv, due to discrimi-
nation against poorer neighborhoods. However, we consider this unlikely.
Caretaker Effects on Aggression and Cortisol Levels
In the current study, at least two aspects of good caretaking may be related to the reduced
aggression and cortisol levels. First, from the interviews with the caretakers, high-level care-
takers spent more time with the cats, before, during, and after feeding, and second, these
caretakers also admitted to being more bonded with the cats they feed, which they expressed
in naming each cat and being able to easily pet and approach most of them. Although the
relationship between free-roaming cats and their caretakers has been previously examined
(Haspel and Calhoon 1990; Natoli et al. 1999), and many caretakers report a strong bond with
the cats they feed (Centonze and Levy 2002), the potentially ameliorating effect of good care-
taking on the welfare of these cats has not been studied. Some relevant ideas on the possi-
ble enrichment effect of good caretaking on free-roaming cats, specifically of time “invested”
in the cats and physical interaction with them, can be derived from studies on other species.
“Low” aggression dogs were reported to be groomed more often by their owners, and own-
ers spent more time with them and had greater attachment to them compared with “high”
aggression dogs (Podberscek 1997). In the case of military dogs, being taken to the handler’s
home (i.e., spending more time with the handler) was associated with improved welfare
(Lefebvre et al. 2007).
Good caretaking was also shown to be related to reduced cortisol levels, indicating lower
stress levels. Other studies have also shown this link: for example, a study on captive clouded
leopards showed that the more time keepers spent with the animals, the lower the animals’
cortisol levels (Wielebnowski et al. 2002). Another study showed that positive human interac-
tion had a moderating effect on the hypothalamic-pituitary-adrenal (HPA) activity of sheltered
dogs (Hennessy et al. 1998; Hennessy et al. 2002). These studies all demonstrate the impor-
tance of positive animal–owner/handler interactions in reducing aggression and improving wel-
fare. Our own findings show that this may also be the case for the cat–human relationship, as
evidenced in the reduced aggression of the cats in the areas with high caretaking. However,
the extent of such influence deserves further attention. If aggression levels can be reduced by
better caretaking, the noise nuisance caused to humans by the cats’ social activity, especially
during the reproductive season (Gunther and Terkel 2002), can be minimized.
Cortisol levels of neutered females were lower at higher caretaking levels. There was also
a tendency for this in intact females, but this was not significant. Females may be more influ-
enced by the caretaker’s actions and behaviors because they roam less compared with males
(Liberg and Sandell 1988) and therefore are more consistently present at the feeding site, and
thus may be in daily contact with the caretaker (Wallace and Levy 2006). Additionally, neutered
females, not needing to search for nesting sites or to have to raise kittens (Bradshaw 1992;
Devillard, Say and Pontier 2003), are expected to roam less than intact females and be more
present at the study site, and thus be more bonded to the caretakers. Consequently, neutered
females can benefit from good caretaking more than other cats in the group, which may be
expressed in reduced cortisol levels, as was previously demonstrated for other species
Finkler et al.
45 Anthrozoös
(Hennessy et al. 1998; Wielebnowski et al. 2002). Another explanation for the reduced cortisol
levels of the neutered females is that of reduction of the excitatory effect of gonadal steroids
on the HPA axis (Seale et al. 2004).
The results of this study demonstrate the need to account for anthropogenic factors when
determining appropriate solutions to cat overpopulation. We recommend that municipal and
private organizations, as well as other individuals involved with cat management and care-
taking, include in their TNR method consideration of the socio-economic status of city areas,
and focus more on those areas that rely mainly on the city’s free-of-charge services, which
according to this study’s results may currently not be sufficient to control the cat populations.
Educational campaigns can help caretakers to learn about the neutering options available
from the municipal veterinary services. High caretaker treatment can contribute to lowering
cats’ aggression levels, thereby both reducing disturbance to people and leading to improved
welfare of the cats. As people in urban areas interact with the cats on a daily basis, it is in our
interest to find new ways to effectively incorporate an understanding of the factors that influ-
ence this interaction into practical cat management, for the benefit of both the human and
the cat populations.
This study was funded in part by the Israel Ministry of Environmental Protection. We thank Ms.
N. Paz for the linguistic editing and Prof. D. Wool, Prof. E. Geffen, Dr. A. Ionescu, and Ms. I.
Gelernter for their statistical assistance. We especially thank Dr. I. Gunther and Dr. E. Shochat
for their critical comments on an earlier version of this manuscript.
Altman, J. 1974. Observational study of behaviour: sampling methods. Behaviour 49: 227–267.
Altschuler, A., Somkin, C. P. and Adler, N. E. 2004. Local services and amenities, neighborhood social capital,
and health. Social Science & Medicine 59(6): 1219–1229.
Alves, M. C. G. P., Matos, M. R. d., Reichmann, M. d. L. and Dominguez, M. H. 2005. Estimation of the dog
and cat population in the State of São Paulo. Revista de Saude Publica 39: 891–897.
Andersen, M. C., Martin, B. J. and Roemer, G. W. 2004. Use of matrix population models to estimate the effi-
cacy of euthanasia versus trap-neuter-return for management of free-roaming cats. Journal of the American
Veterinary Medical Association 225(12): 1871–1876.
Azienda USL Roma D. <> Accessed November 2009.
Baker, P. J. and Harris, S. 2007. Urban mammals: What does the future hold? An analysis of the factors affecting
patterns of use of residential gardens in Great Britain. Mammal Review 37(4): 297–315.
Beck, A. M. 2002. The Ecology of Stray Dogs: A Study of Free-Ranging Urban Animals. West Lafayette, IN: Pur-
due University Press.
Bradshaw, J. W. S. 1992. The cat–human relationship. In The Behaviour of the Domestic Cat, 163–176. Wolling-
ford, UK: CAB International.
Bradshaw, J. W. S., Horsfield, G. F., Allen, J. A. and Robinson, I. H. 1999. Feral cats: Their role in the popula-
tion dynamics of Felis catus. Applied Animal Behaviour Science 65(3): 273–283.
Calhoon, R. E. and Haspel, C. 1989. Urban cat populations compared by season, subhabitat and supplemental
feeding. Journal of Animal Ecology 58(1): 321–328.
Centonze, L. A. and Levy, J. K. 2002. Characteristics of free-roaming cats and their caretakers. Journal of the
American Veterinary Medical Association 220(11): 1627–1633.
Childs, J. E. 1990. Urban cats: Their demography, population density, and owner characteristics in Baltimore,
Maryland. Anthrozoös 3: 234–244.
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
46 Anthrozoös
Christensen, T. and Leaegreid, P. 2002. Trust in Government—The Relative Importance of Service Satisfaction,
Political Factors and Demography. Stein Rokkan Centre for Social Studies, Bergen University Research
Church, S. C., Allen, J. A. and Bradshaw, J. W. S. 1996. Frequency-dependent food selection by domestic cats:
A comparative study. Ethology 102(6): 495–509.
Clarke, A. L. and Pacin, T. 2002. Domestic cat “colonies” in natural areas: A growing exotic species threat.
Natural Areas Journal 22(2): 154–159.
Cohen, S., Doyle, W. J. and Baum, A. 2006. Socioeconomic status is associated with stress hormones.
Psychosomatic Medicine 68(3): 414–420.
Dards, J. L. 1983. The behaviour of dockyard cats: Interactions of adult males. Journal of Applied Ethology 10:
Delhey, J. and Newton, K. 2002. Who Trusts? The Origins of Social Trust in Seven Nations. Social Science Re-
search Center Berlin (WZB), Research Unit “Social Structure and Social Reporting.”
Devillard, S., Say, L. and Pontier, D. 2003. Dispersal pattern of domestic cats (Felis catus) in a promiscuous urban
population: Do females disperse or die? Journal of Animal Ecology 72(2): 203–211.
de Walle, H. E. K. and de Jong-van den Berg, L. T. W. 2007. Growing gap in folic acid intake with respect to
level of education in the Netherlands. Community Genetics 10(2): 93–96.
Finkler, H. and Terkel, J. 2010. Cortisol levels and aggression in neutered and intact free-roaming female cats
living in urban social groups. Physiology & Behavior 99(3): 343–347.
Finkler, H. and Terkel, J. in press. Dichotomy in the emotional approaches of caretakers of free-roaming cats in
urban feeding groups: findings from in-depth interviews. Anthrozoös.
Gdalevich, M., Mimouni, D., Ashkenazi, I. and Shemer, J. 2000. Rabies in Israel: Decades of prevention and a
human case. Public Health 114(6): 484–487.
Gunther, I. and Terkel, J. 2002. Regulation of free-roaming cat (Felis silvestris catus) populations: A survey of the
literature and its application to Israel. Animal Welfare 11(2): 171–188.
Haspel, C. and Calhoon, R. E. 1990. The interdependence of humans and free-ranging cats in Brooklyn, New
York. Anthrozoös 3: 155–161.
Haspel, C. and Calhoon, R. E. 1993. Activity patterns of free-ranging cats in Brooklyn, New-York. Journal of
Mammalogy 74(1): 1–8.
Hennessy, M. B., Voith, V. L., Hawke, J. L., Young, T. L., Centrone, J., McDowell, A. L., Linden, F. and Daven-
port, G. M. 2002. Effects of a program of human interaction and alterations in diet composition on activity of
the hypothalamic-pituitary-adrenal axis in dogs housed in a public animal shelter. Journal of the American
Veterinary Medical Association 221(1): 65–71.
Hennessy, M. B., Williams, T., Miller, D. D., Douglas, C. W. and Voith, V. L. 1998. Influence of male and female
petters on plasma cortisol and behaviour: Can human interaction reduce the stress of dogs in a public ani-
mal shelter? Applied Animal Behaviour Science 61(1): 63–77.
Hughes, K. L. and Slater, M. R. 2002. Implementation of a feral cat management program on a university cam-
pus. Journal of Applied Animal Welfare Science 5(1): 15–28.
Israeli Central Bureau of Statistics 2008. Statistical abstract of Israel 2008: Localities, population and density per
sq km, by metropolitan area and selected localities. <
shnatone_new.htm?CYear=2008&Vol=59&CSubject=2> Accessed November 2009.
ISTAT- Instituto Nazionale Di Statistica 2005. 14 Censimento generale della popolazione e delle abitazioni 2001.
<> Accessed November 2009.
Izawa, M. 1983. Daily activities of the feral cat Felis catus LINN. The Journal of Mammalogical Society of Japan
9(5): 219–228.
Jessup, D. A. 2004. The welfare of feral cats and wildlife. Journal of the American Veterinary Medical Association
225(9): 1377–1383.
Kinzig, A., Warren, P. S., Martin, C., Hope, D. and Madhusudan, K. 2005. The effects of human socioeconomic
status and cultural characteristics on urban patterns of biodiversity. Ecology and Society 10(1): 23.
Lefebvre, D., Diederich, C., Delcourt, M. and Giffroy, J.-M. 2007. The quality of the relation between handler and
military dogs influences efficiency and welfare of dogs. Applied Animal Behaviour Science 104(1–2): 49–60.
Levy, J. K. and Crawford, P. C. 2004. Humane strategies for controlling feral cat populations. Journal of the
American Veterinary Medical Association 225(9): 1354–1360.
Finkler et al.
47 Anthrozoös
Levy, J. K., Gale, D. W. and Gale, L. A. 2003. Evaluation of the effect of a long-term trap-neuter-return and
adoption program on a free-roaming cat population. Journal of the American Veterinary Medical Association
222(1): 42–46.
Liberg, O. and Sandell, M. 1988. Spatial organization and reproductive tactics in the domestic cat and other
felids. In The Domestic Cat: Tthe Biology of its Behaviour, 83–98, ed. D. C. Turner and P. Bateson. Cambridge:
Cambridge University Press.
Markovchick-Nicholls, L., Regan, H. M., Deutschman, D. H., Widyanata, A., Martin, B., Noreke, L. and Hunt,
T. A. 2008. Relationships between human disturbance and wildlife land use in urban habitat fragments.
Conservation Biology 22(1): 99–109.
Mirmovitch, V. 1995. Spatial organization of urban feral cats (Felis Catus) in Jerusalem. Wildlife Research 22(3):
Morley, B., Chapman, K., Mehta, K., King, L., Swinburn, B. and Wakefield, M. 2008. Parental awareness and
attitudes about food advertising to children on Australian television. Australian and New Zealand Journal of
Public Health 32(4): 341–347.
Natoli, E. 1985. Spacing pattern in a colony of urban stray cats (Felis catus L.) in the historic centre of Rome.
Applied Animal Behaviour Science 14(3): 289–304.
Natoli, E. 1994. Urban feral cats (Felis catus L.): Perspectives for a demographic control respecting the psycho-
biological welfare of the species. Annali dell Istituto Superiore Di Sanita (Roma) 30(2): 223–227.
Natoli, E., De Vito, E. and Pontier, D. 2000. Mate choice in the domestic cat (Felis silvestris catus L.). Aggressive
Behavior 26(6): 455–465.
Natoli, E., Ferrari, M., Bolletti, E. and Pontier, D. 1999. Relationships between cat lovers and feral cats in Rome.
Anthrozoös 12(1): 16–23.
Natoli, E., Maragliano, L., Cariola, G., Faini, A., Bonanni, R., Cafazzo, S. and Fantini, C. 2006. Management
of feral domestic cats in the urban environment of Rome (Italy). Preventive Veterinary Medicine 77(3–4): 180–
Nutter, F. B., Levine, J. F. and Stoskopf, M. K. 2004. Reproductive capacity of free-roaming domestic cats and
kitten survival rate. Journal of the American Veterinary Medical Association 225(9): 1399–1402.
Oppenheimer, E. 1980. Felis catus population densities in urban areas. Carnivore Genetic Newsletter 4: 72–80.
Podberscek, A. L. 1997. Illuminating issues of companion animal welfare through research into human–animal
interactions. Animal Welfare 6(4): 365–372.
Porat, I., Frenkel, A. and Shoshany, M. 2008. Residential and nonresidential construction initiations in Tel Aviv-
Yafo: Autocorrelation analysis of urban structure evolution. Environment and Planning B-Planning & Design
35(3): 535–551.
Rees, P. 1981. The ecological distribution of feral cats and the effects of neutering a hospital colony Proceed-
ings of a Symposium on the Ecology and Control of Feral Cats, 12–22. Potters Bar, Hertfordshire: Universi-
ties Federation for Animal Welfare.
Remfry, J. 1981. Strategies for control. Proceedings of a Symposium on the Ecology and Control of Feral Cats,
73–80. Potters Bar, Hertfordshire: Universities Federation for Animal Welfare.
Schmidt, P. M., Lopez, R. R. and Collier, B. A. 2007. Survival, fecundity, and movements of free-roaming cats.
The Journal of Wildlife Management 71(3): 915–919.
Seale, J. V., Wood, S. A., Atkinson, H. C., Bate, E., Lightman, S. L., Ingram, C. D., Jessop, D. S. and Harbuz,
M. S. 2004. Gonadectomy reverses the sexually diergic patterns of circadian and stress-induced hypothal-
amic-pituitary-adrenal axis activity in male and female rats. Journal of Neuroendocrinology 16(6): 516–524.
Shaw, L. M., Chamberlain, D. and Evans, M. 2008. The House Sparrow Passer domesticus in urban areas:
Reviewing a possible link between post-decline distribution and human socioeconomic status. Journal of
Ornithology 149(3): 293–299.
Slater, M. R. 2001. The role of veterinary epidemiology in the study of free-roaming dogs and cats. Preventive
Veterinary Medicine 48(4): 273–286.
Strula, K. 1993. Role of breeding regulation laws in solving the dog and cat overpopulation problem. Journal of
the American Veterinary Medical Association 18 (773): 794.
Toukhsati, S. R., Bennett, P. C. and Coleman, G. J. 2007. Behaviors and attitudes towards semi-owned cats.
Anthrozoös 20: 131–142.
Wallace, J. L. and Levy, J. K. 2006. Population characteristics of feral cats admitted to seven trap-neuter-
return programs in the United States. Journal of Feline Medicine & Surgery 8(4): 279–284.
The Impact of Anthropogenic Factors on the Behavior, Reproduction…
48 Anthrozoös
Wielebnowski, N. C., Fletchall, N., Carlstead, K., Busso, J. M. and Brown, J. L. 2002. Noninvasive assessment
of adrenal activity associated with husbandry and behavioral factors in the North American clouded leopard
population. Zoo Biology 21(1): 77–98.
Williams, D. R. 1998. Race, socioeconomic status, and health the added effects of racism and discrimination.
Annals New York Academy of Sciences 896: 173–188.
Xemar, V., Pontier, D. and Artois, M. 1998. Attempt to control stray cats in an urban area. Paper presented at
the 8th International Conference on Human–Animal Interactions “The Changing Role of Animals in Society,”
Prague, Czech Republic, September 10–12, 1998.
Finkler et al.
49 Anthrozoös
... Groups of FRCs are also common in city centers and residential areas. Research has included FRCs living at locations such as Regent's Park in London, England [32], in urban and residential areas of Israel [33][34][35][36], and in the urban centers of Rome, Italy. These locations include the work of Natoli and colleagues in the market square "Piazza Vittorio Emanuele" [37], a large courtyard known as "Garbatella" [38][39][40] (which had a density of 2833 cats per km 2 [41]), and the historical ruins near the center of Rome, "Fori di Traiano" [42]. ...
... In one study, aggression was rare both before and following neutering [32]. In other work, neutered groups of cats displayed less aggression than unneutered groups of cats [33] and compared to intact cats, neutered cats showed lower frequencies of agonistic behavior [35]. Aggressive behaviors have also been compared with cortisol [34]. ...
... Finally, level of human caretaking may also impact aggressive behavior. In one study, cats who had minimal human care displayed higher aggression than cats who received maximum care [35]. In all, human activities such as sterilization, provisioning of food, and level of care, as well as traits of conspecifics (such as their age or relationship to one another) are all important factors impacting aggressive behavior to varying degrees ( Table 2). ...
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Despite the diversity of social situations in which cats live, the degree to which free-ranging cats (FRCs) are social is still debated. The aim of this review is to explore the literature on the social behavior of FRCs. A search of two major databases revealed that observations of intraspecies and interspecies social interactions have been conducted. The intraspecific social dynamics of FRCs differ based on group of cats surveyed. Some groups display strong social bonds and preferential affiliations, while other groups are more loosely associated and display little to no social interaction. Factors impacting FRC conspecific interactions include cat body size, cat social rank, cat individuality, cat age, relationship to conspecific (kin/familiar), cat sex, level of human caretaking, presence of food, the health of the individual, or sexual status of conspecifics. Interspecies interactions also occur with humans and wildlife. The human’s sex and the weather conditions on the day of interaction have been shown to impact FRC social behavior. Interactions with wildlife were strongly linked to the timing of cat feeding events. These findings support the idea that FRCs are “social generalists” who display flexibility in their social behavior. The social lives of FRCs exist, are complex, and deserve further study.
... Cat management decisions to preserve biodiversity would benefit from considering the spatial location of outdoor feeding sites. Currently, other municipalities may find it difficult to enact these management recommendations since feeding locations are often run by individuals and their locations are not readily known (Finkler et al., 2011). We recommend that stakeholders work together to document and share data on free-roaming cat feeding locations, while ensuring the privacy of individuals who feed cats. ...
... The ecological loss inflicted by cats is well-documented (Loss et al., 2013). While some non-native species are relatively selfsufficient and are largely beyond human control, cats pose a unique situation in which their continued persistence on the landscape can be largely attributed to human actions (Doherty et al., 2014;Lepczyk et al., 2011;Finkler et al., 2011). Professionals across relevant fields largely agree that cats face and impose risks when outdoors, however proposed management actions differ (Foreman-Worsley et al., 2021;Leong et al., 2020;Sherwood et al., 2019;Wald et al., 2013). ...
The ecological impact of free-roaming domestic cats (Felis catus) is well-studied. However, despite receiving considerable attention in both the scientific and popular literature, predation behavior is rarely an explicit consideration when developing cat population management plans. We used motion-activated wildlife cameras to document predation events by cats in Washington, D.C. (U.S.A), and assessed the relationships between predation and local environmental characteristics. Our analyses reveal that predation by cats is greatest where supplemental food is most abundant, and that the probability of a cat preying upon a native species increases closer to forest edges. Conversely, we found that the probability of a cat depredating a non-native brown rat increases with increasing distance from forest edges. Therefore, we recommend the implementation of cat exclusionary buffer zones around urban forests and that free-roaming domestic cat management policies explicitly consider the spatial location of cat-feeding sites. Our findings provide a data-driven approach to free-roaming cat management.
... This finding also supports caution regarding neutering and returning cats. While some studies show improved welfare of TNR cats based on reductions in fighting [63,76], improved longevity [62], lower rates of infectious disease [6], and healthier appearance [63], we share the concerns of some animal welfare groups, such as People for the Ethical Treatment of Animals (PETA), that returned cats may not receive long-term caregiver support and are still highly vulnerable to trauma [77,78]. ...
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Globally, unowned urban cats are a major concern because they may suffer from poor welfare and cause problems, including public health risks, nuisances, and urban wildlife predation. While management options are often presented as a choice between culling or trap–neuter–return (TNR), for 25 years, the Lonely Miaow (Inc.) charity in Auckland, New Zealand (hereafter LM), has used a third strategy—intensive adoption or trap–assess–resolve (TAR). As of 2019, of 14,611 unowned cats trapped, 64.2% were adopted, 22.2% were euthanized if unsocialised or in grave ill-health, 5.7% were neutered and returned to the site, and 7.9% had other outcomes, such as being transferred to other shelters. Adoption rates increased over this time, exceeding 80.0% in 2018 and 2019. The cost of processing each cat from capture to adoption rose from NZD 58 in 1999 to NZD 234 by 2017. Approximately 80% of colonies (sites where cats were trapped) were around residential areas. Approximately 22% of cats required veterinary treatment after capture; common ailments included respiratory infections, ringworm, dental problems, and trauma. Consistently, 52% of cats were young kittens (<10 weeks old), c. 80% of cats were <1 year old, and only c. 2% were estimated to be >5 years old. TAR avoids euthanasia where possible. Its effectiveness would be enhanced by fewer abandonments of owned cats and kittens, fitting within integrated strategies for the control of unowned cats involving community education. Cat adoptions improve the welfare of cats and, with appropriate husbandry, should alleviate concerns about nuisances, public health, and attacks on wildlife or the cats themselves, essentially benefitting the community and the cats. This case study is relevant to other cities around the world that are seeking to manage unowned cats.
... Further work in this area is warranted especially as many cat owners hold misconceptions regarding the age of puberty of cats and are unaware that related cats can mate with each other 36 , which is a particular concern when owners acquire littermates when obtaining new kittens. Additionally, we find strong links between human deprivation and unowned cats, likely mediated by lower rates of neutering of owned and unowned cats within these communities 37,38 . This outcome is likely related to a balance of multiple socio-economic factors and their associated barriers influencing capability, opportunity and motivation 39 to neuter cats. ...
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Globally, unowned cats are a common element of urban landscapes, and the focus of diverse fields of study due to welfare, conservation and public health concerns. However, their abundance and distribution are poorly understood at large spatial scales. Here, we use an Integrated Abundance Model to counter biases that are inherent in public records of unowned cat sightings to assess important drivers of their abundance from 162 sites across five urban towns and cities in England. We demonstrate that deprivation indices and human population densities contribute to the number of unowned cats. We provide the first spatially explicit estimates of expected distributions and abundance of unowned cats across a national scale and estimate the total UK urban unowned cat population to be 247,429 (95% credible interval: 157,153 to 365,793). Our results provide a new baseline and approach for studies on unowned cats and links to the importance of human-mediated effects.
... Taking into account the characteristics of the inhabitants and the habitat of the area of residence is complex but crucial for effective stray cat policy (Lord, 2008;Mameno et al., 2017). All these variables play a role in designing policies, targeting communication and creating public awareness (Deak et al., 2019;Finkler et al., 2011b;Flockhart et al., 2016). As Crowley et al. (2020) states: the dual status of the species as both wild predator and domestic companion underpins different interaction with and nuisance for humans. ...
Stray cat population management is an important worldwide issue. Understanding citizen attitudes towards stray cat control options is vital to the success of controlling stray cat numbers, as public perception affects the acceptance of, support for and collaboration in stray cat management policies. Audience segmentation, as to enable each group to be engaged in the stray cat management policy, is important for the success of the interventions. Therefore a web-based survey was conducted among Flemish citizens in order to examine differences in acceptance towards seven management scenarios: household cat neutering with financial support for the owner, household cat neutering without financial support for the owner, encouraging responsible household cat ownership, trapping stray cats and taking them to a shelter, trapping and neutering stray cats for release into a managed “cat colony” (composed by so called “community cats”), trapping and killing of stray cats, and undertaking no action. A total of 4059 valid responses were collected and the proportions of agreement were compared across the different management scenarios using the two-sample z-test. Interactions among factors that influenced each management scenario were investigated using the CHAID (Chi-squared Automatic Interaction Detection) analysis and visualized on a tree. Our results showed that fostering responsible household cat ownership (89.9%) and conversion of stray cats to “community cats” (76.3%) were most supported by respondents in our sample (which consisted mainly of females, cat-lovers, and families without children). Least supported were the killing of stray cats (7.7%) and undertaking no action (3.3%). The demographic analyses revealed that for the acceptance of management scenarios there were three important factors (attitude towards cats, area of residence, and gender), two weaker factors (education and having children) and two which had almost no impact (age and cat ownership). We propose that future studies should focus on the effect of ‘area of residence’, ‘having children’ and ‘education’. In conclusion, our research confirms that management of and communication on stray cat strategies should not be developed with a one-size-fits-all approach. Efforts should be tailored to each audience segment, thus adapted to the area of residence and human characteristics.
... 11 Although more work is needed to explore the dynamic nature and fluidity of cat populations, consistency in messaging across the veterinary profession is key to ensure owners and cats are provided with the same standard of care, even if locally the problem is difficult to conceptualise. It is anticipated that the neutering rates of unowned cats are lower 42 and their abundance higher 3 43 in areas of social deprivation; however, we found no links between IMD and prepubertal neutering. This result indicates that there are veterinarians across deprived and affluent areas carrying out the procedure, and that not all veterinarians hold the previously mentioned view of some VNOFs that it is not applicable in some areas. ...
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Background Many cats in the UK are not neutered before reaching potential breeding age. The purpose of this investigation was to understand the prevalence of veterinarians’ behaviours around neutering cats at four months of age and to identify any needs of the veterinary profession on this. Methods A sample of 483 veterinarians that completed a cross-sectional online questionnaire were included in quantitative and qualitative analyses. Results Almost 70 per cent of veterinarians indicated they were comfortable carrying out neutering on cats of four months of age, and approximately half of veterinarians indicated they would recommend neutering client-owned cats at four months of age if practice policy permitted. There was no association found between these practices and geographic risk factors. Instead, neutering at four months was associated with the gender of the veterinarian, their practice policy and whether they routinely neuter unowned cats. Veterinarians have contrasting beliefs on similar themes, depending on whether they neuter kittens at four months or not, including differing opinions on general anaesthetic, surgery risks, owner compliance and their perceptions of neutering practices within the wider profession. Conclusion Familiarity and experience with the procedure, alongside perceived norms, are central to beliefs. We also highlight informational and learning barriers.
... The link between felines and T. gondii is well documented (Suzán and Ceballos 2005;Vanwormer et al. 2013;Barros et al. 2018), and the size and spatial distribution of outdoor cat populations are closely tied to local human populations (Ferreira et al. 2011;Finkler et al. 2015). Unmanaged outdoor cat populations are known to shed large numbers of oocysts that are distributed across the landscape via stormwater runoff (Miller et al. 2002(Miller et al. , 2008 and infect native wild species (e.g., otters; Vanwormer et al. 2013). ...
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The North American river otter ( Lontra canadensis) is the largest mustelid in North Carolina, USA, and was once extirpated from the central and western portions of the state. Over time and after a successful reintroduction project, otters are now abundant and occur throughout North Carolina. However, there is a concern that diseases may have an impact on the otter population, as well as on other aquatic mammals, either through exposure to emerging diseases, contact with domestic animals such as domestic cats ( Felis catus), or less robust condition of individuals through declines in water quality. We tested brain and kidney tissue from harvested otters for the pathogens that cause leptospirosis, parvovirus, and toxoplasmosis. Leptospirosis and toxoplasmosis are priority zoonoses and are maintained by domestic and wild mammals. Although parvovirus is not zoonotic, it does affect pets, causing mild to fatal symptoms. Across the 2014–15 and 2015–16 trapping seasons, we tested 220 otters (76 females, 144 males) using real-time PCR for Leptospira interrogans, parvovirus, and Toxoplasma gondii. Of the otters tested, 1% (3/220) were positive for L. interrogans, 19% (41/220) were positive for parvovirus, and 24% (53/220) were positive for T. gondii. Although the pathogens for parvovirus and toxoplasmosis are relatively common in North Carolina otters, the otter harvest has remained steady and the population appears to be abundant and self-sustaining. Therefore, parvovirus and toxoplasmosis do not currently appear to be negatively impacting the population. However, subsequent research should examine transmission parameters between domestic and wild species and the sublethal effects of infection.
... Seo and Tanida [18] reported that health conditions were judged to be bad in about half of feral cats living in the uptown and downtown areas, and it was presumed that most missing cats died from illness or injury, as opposed to having migrated elsewhere in the west of Japan. Comparing the groups of stray cats with and without a high level of human care in the city area of Tel Aviv, Israel, stray cats receiving a high level of care showed low aggressiveness and low fecal cortisol concentration in spayed females [19]. It turned out that stray cats are influenced by people, and that, even if they were stray, the involvement of people with care leads to the improvement in the welfare of cats. ...
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The community cats program (CCP), which includes trap–neuter–return activities, has been promoted in Japan to reduce the population of free-roaming cats without harmful effects on their welfare. To ascertain the effects of the CCP, a two-year route census of free-roaming cats was conducted in an area with CCP and the other area without CCP in urban Tokyo, Japan. The estimated number of cats was lower in the CCP area than the non-CCP area, but there was no difference in the rate of decline in cat populations between areas. More cats emigrated or disappeared rather than immigrated in both areas in the second year and more males tended to immigrate into the CCP area. There was no difference in the behavior of cats between areas and among seasons. The proportion of cats with poor health was lower in the CCP area than the non-CCP area. These results suggest that the CCP may improve the welfare of free-roaming cats. As the effect of CCP was restrictive in reducing the population of free-roaming cats, the further promotion of neutering of cats may be necessary to reduce the population density of cats.
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Trap-neuter-return (TNR) is promoted as a "humane" alternative to lethal methods for population control of feral domestic cats (Felis catus). This paper explores feed-backs between feral domestic cats, coyotes (Canis latrans), raccoons (Procyon lotor), and skunks (Mephitis mephitis) at a TNR feral cat colony in Rhode Island, USA. A total of 12,272 photographs from a motion-activated camera were analyzed. Cat population size and visitation frequency of wildlife were estimated during three different feeding regimes. Abundant food on the ground was associated with increased wildlife visits, while elevated or limited food was associated with decreased wildlife visits. During the two-year study period, the population of cats dropped from 17 to 12 individuals and
Feces littered on the ground by free-roaming cats contain bacteria, viruses, and parasites and pose a significant health risk to humans. The purpose of this study was to examine the effect of communal litter box provision on the defecation behavior of a free-roaming cat population. The study was conducted at H temple and its graveyard in the uptown area of old-town Onomichi, Japan. Cat feces were collected and weighed once a week for 4 weeks, at five popular defecation sites in the temple precincts and graveyard, to assess the quantity of feces left by the cats. A commercial cat repellent was then applied to the ground at 11 sites, including the five popular defecation sites, and six communal litter boxes, created by filling repurposed plastic planters with cat litter, were provided at different sites. The feces in the six litter boxes and on the ground at the five defecation sites were collected and weighed once a week for 14 weeks. The behavior of the cats around the litter boxes and defecation sites was captured using trail cameras. The total weight of the feces collected from the ground before the application of the litter boxes and cat repellent was 939 g. Three adult cats were mainly responsible for the feces on the ground. The amount of feces found on the ground around the temple decreased gradually and significantly after the provision application of the litter boxes and repellent, and reached 0 g in the final week of the study. In contrast, the average weight of the feces in the six litter boxes increased gradually and significantly, and reached 65.7 g/litter box/week in the 14th week. The results showed that the provision of litter boxes and the use of repellent is effective in changing the defecation behavior of ownerless free-roaming cats. We recommend promoting the provision of litter boxes to free-roaming cats to reduce fecal pollution in Onomichi and engaging with local cat feeders to participate in the management of the litter boxes, such as cleaning and changing the litter.
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Although the occurrence of cat-caretaking of free-roaming cats is widespread, particularly so in countries with a climate suitable for cats to reproduce year-round, our knowledge of this relationship is still incomplete. People who engage in daily activities of feeding and caring for groups of free-roaming cats (cat caretakers) are known to be devoted to their cats and invest considerable resources in their care, including neutering and veterinary care. These caretakers often encounter difficulties, such as resentment by neighbors and lack of cooperation or financing by the municipal veterinary services. Despite the fundamental understanding of these caretakers' high daily commitment, and sometimes strong bond with the cats, detailed knowledge is still lacking regarding the nature of this bond, the difficulties that ensue from this daily occupation, and the relationship between the two. The purpose of this study was thus to acquire a deeper understanding, by means of an in-depth interview with cat caretakers. The study has identified, for the first time, two distinct emotional approaches that accompany extensive caretaking for free-roaming cats: emotional attachment and emotional detachment. We show how these two different responses affect both social and financial aspects in the caretakers' lives, and report on the ways in which these individuals experience cat caretaking. Our findings provide a first systematic understanding of the relationship between the level of technical caretaking (feeding, medical care, etc.) and the level of emotional involvement, and reveal the ambivalence often inherent in human–animal relations in general and the caretaker–cat bond in particular. The understanding acquired here can be put into practice to reduce the emotional and technical difficulties experienced by cat caretakers, as well as to improve free-roaming cat management efforts and cat welfare. By increasing public and municipal awareness of the possible contribution of cat caretakers to cat management, and of the emotional and technical difficulties they experience, both the caretakers and other community members can benefit.
The second edition of this book contains 12 chapters that discuss the domestication and general biology (anatomy and physiology), senses, behaviour mechanisms, behavioural development, communication, feeding behaviour, hunting and predatory behaviours, social behaviour, relationship with humans, welfare, abnormal/unwanted behaviour and the physiological and pathological causes of behavioural changes. The book contains some tables, black and white illustrations, a list of references for further reading and an index. It will be of use to those with an interest in cat behaviour.
Preferences for common food types ('apostatic selection') have been demonstrated in a wide variety of vertebrate predators, yet there are few examples of preferences for rare food types ('anti-apostatic selection'). Anti-apostatic selection is predicted to occur when, among other things, there are nutritional benefits to be gained from the consumption of a mixed diet. We tested this hypothesis by examining the frequency- dependent food preferences of domestic cats (Felis silvestris catus) with different nutritional histories. Subjects were classified as being either nutritionally 'experienced' (farm and rescue shelter cats, with a history of scavenging for nutritionally variable foods) or nutritionally 'inexperienced' (cats reared indoors on high-quality, nutritionally complete diets). We tested for frequency dependence by allowing individuals and groups of cats from the two groups to select from high-density mixtures of two types of artificial food pellet. In experiments on individual cats, nutritionally experienced subjects showed significant anti-apostatic selection, whereas inexperienced cats produced only a weak anti apostatic trend. In experiments on groups of cats, both inexperienced and experienced groups showed significant anti- apostatic selection. The apparent inconsistency between individual and group results could be explained in terms of the additional anti apostatic effects that result from variation among individuals in group foraging situations (i.e. when the effects of individuals are pooled). Because other behavioural explanations, such as perceptual contrast and sampling effects, were unlikely to have influenced our results, we conclude that the differences in selection between experienced and inexperienced individuals were probably due to the differing extent to which the consumption of a mixed diet was beneficial. These experiments may offer some insight into the success of the domestic cat in urban areas: although obligate carnivores, they appear to possess flexible feeding strategies which will tend to allow them to select a reasonably balanced diet from nutritionally variable resources in, for example, refuse bins.
The domestic cat (Felis catus L.) population in the United States has more than doubled since 1970 and is estimated to exceed 100 million animals. Domestic cats are considered a direct predation threat to native wildlife, and, in a growing number of U.S. cities, conflicts are arising between land managers responsible for conserving native wildlife and members of the public concerned with the welfare of feral and abandoned cats. The removal of cats from natural areas has a strong sociopolitical component, including the support of a well-organized, well-funded special interest citizen group prepared to resist removal of cats. In addition, land managers must address removal within the larger political framework in which their agency is embedded. We describe the increasingly common trap-and-release approach to feral and abandoned cat management employed by cat welfare organizations. We use as examples two colony management organizations operating in south Florida. Land managers should take a proactive approach to the issue of feral and abandoned cats and undertake review of existing ordinances prior to the establishment of a significant cat population. Conservationists must expand their efforts to include support for the long-term effort to educate the public on the importance of responsible pet ownership and the establishment and enforcement of appropriate pet ordinances.
Population densities of free-ranging Felis catus were compared in 2 contiguous urban subhabitats. One subhabitat, characterized by voluminous, poorly contained refuse, and many abandoned buildings, supported 4.88 ± 0.82 cats ha-1, which differed significantly from the 2.03 ± 0.2 cats ha-1 supported by the other subhabitat (partial refuse containment, few abandoned buildings). Neither season nor supplemental feeding had a significant effect on population density. The distribution of individuals within the study area varied with availability of shelter and was not dependent upon food. -from Authors
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
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.
Feral cats were studied for 10 months in a residential area in Jerusalem and their spatial distribution compared during two 1-month periods, the first in the autumn prior to the mating season and the second during the mating season (winter). Cat locations were recorded by direct observations, and home-range sizes were calculated with the minimum convex polygon method. No significant change in home-range size of adult males or females was found between the 2 periods. Young males expanded their home ranges considerably during their first mating season. Home ranges of males were significantly larger than those of females in both periods (0.56 and 0.30 ha, respectively, in autumn; 0.75 and 0.27 ha in winter). The home ranges of both sexes overlapped considerably with individuals of the same sex. Overlap among home ranges of females indicated a group pattern. High overlap (80%) was found among females that fed from the same set of garbage bins with similar frequency. Lower overlap (20%) was found between individual females that shared only a subset of their food resources and used it with different frequency. It is suggested that the distribution of food patches (garbage bins), the amount of food available and the rate of food renewal determined the cats' spatial organisation.