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Citation: Calver, M.C.; Crawford,
H.M.; Scarff, F.R.; Bradley, J.S.;
Dormon, P.; Boston, S.; Fleming, P.A.
Intensive Adoption as a Management
Strategy for Unowned, Urban Cats:
A Case Study of 25 Years of
Trap–Assess–Resolve (TAR) in
Auckland, New Zealand. Animals
2022,12, 2301. https://doi.org/
10.3390/ani12172301
Academic Editor: Mandy Paterson
Received: 8 August 2022
Accepted: 29 August 2022
Published: 5 September 2022
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animals
Article
Intensive Adoption as a Management Strategy for Unowned,
Urban Cats: A Case Study of 25 Years of Trap–Assess–Resolve
(TAR) in Auckland, New Zealand
Michael C. Calver 1, *, Heather M. Crawford 1, Fiona R. Scarff 1, J. Stuart Bradley 1, Peter Dormon 2,
Samantha Boston 2and Patricia A. Fleming 3
1Environmental and Conservation Sciences, Murdoch University, 90 South Street,
Murdoch, WA 6150, Australia
2Lonely Miaow Association, Lonely Miaow Association, 35a, Riddell Road, Glendowie,
Auckland 1071, New Zealand
3Harry Butler Institute, Murdoch University, 90 South Street, Murdoch, WA 6150, Australia
*Correspondence: m.calver@murdoch.edu.au
Simple Summary:
Unowned urban cats may suffer from poor welfare and cause problems, e.g.,
public health risks, nuisances, and urban wildlife predation. For 25 years, the Lonely Miaow (Inc.)
charity in Auckland, New Zealand, has used intensive adoption to rehome unowned cats. By the end
of 2019, LM volunteers had trapped 14,611 unowned cats, which were adopted wherever possible
(64.2%), euthanized if unsocialised or in grave ill-health (22.2%), or (infrequently) neutered and
returned to the site (5.7%). The remaining 7.9% had other outcomes, such as being transferred to other
shelters. Adoption rates increased over 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 in 2017. Approximately 80%
of colonies (sites where cats were trapped) were around residential areas. Most cats were young and
very few were over 5 years old. Around one in five cats needed veterinary treatment, with respiratory
infections and injury common. Adopting cats and removing them from the streets improves their
welfare, essentially benefitting the community and the cats. The effectiveness of adoption strategies
would be enhanced by fewer abandonments of owned cats and kittens, fitting within integrated
strategies for the control of unowned cats involving community education.
Abstract:
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.
Keywords: TNR; free-roaming cat; stray cat; euthanasia; cat welfare; adoption
Animals 2022,12, 2301. https://doi.org/10.3390/ani12172301 https://www.mdpi.com/journal/animals
Animals 2022,12, 2301 2 of 21
1. Introduction
Domestic cats, Felis catus, are prolific breeders, producing litters of up to four kittens
two to three times a year [
1
], (Chapter 4). Thus, one unfortunate consequence of their
global popularity as companion animals [
2
,
3
] is the potential for large populations of
unowned, free-ranging cats around human habitations, sustained by the abandonment of
owned animals and uncontrolled breeding [
4
–
8
]. They are variously named unowned cats,
free-ranging cats, stray cats, community cats, or feral cats, with little agreement on a chaotic
nomenclature [
9
,
10
]; see Figure 1for the nomenclature that we use herein. The numbers of
unowned cats are difficult to quantify, with estimates of their abundance ranging between
30 and 100 million in the USA (Ref. [
11
] and included references), 700,000 in Australia [
12
],
and 196,000 in New Zealand [
13
]. Depending on the geographic region, stray cats may
suffer from similar poor welfare outcomes to free-ranging owned cats, including road
accident trauma [
14
–
17
], poisoning [
18
], ingestion of hazardous garbage [
7
], predation by
larger carnivores [
19
], disease [
20
], and human persecution [
21
]. Free-ranging cats may also
kill or harass wildlife [
22
–
24
], including threatened species [
25
–
27
], present a disease risk
to people, pets, and wildlife [
28
–
32
], and annoy people [
1
,
33
]. Therefore, the management
of stray cats is an ethical and community concern.
Figure 1.
Categories of cats under definitions provided by the New Zealand Government’s Animal
Welfare (Companion Cats) Code of Welfare 2018. Sexually entire cats may interbreed across categories.
While the scale of the problem of stray cat management requires intervention, there
is an extensive debate regarding the most effective and ethical options (e.g., contrast [
34
]
with [
1
], or [
35
–
37
] with [
8
,
38
]). Options are often constrained by legislation, which varies
greatly between jurisdictions [
39
–
43
]. Trapping and euthanizing cats (TE) [
37
,
44
] sometimes
meets with community resistance. There has been a growing number of literature reports
describing alternative actions for potentially resolving stray cat issues, especially regarding
trapping, neutering, and returning cats to the site of capture, e.g., trap–neuter–return (TNR);
Animals 2022,12, 2301 3 of 21
with variants such as TNVR (where cats are vaccinated before release) or TTNR (where
there is intense targeting of a restricted geographical area), e.g., [
45
,
46
]. These methods
entail varying degrees of follow-up support, including substantial time and money invested
by ‘caretakers’ for these cats. There are strong advocates and opponents of TNR in all its
options [8,35–38].
New Zealand has very high cat ownership rates, with c. 35% of households owning
one or more cats [
47
]. In common with many other countries, the popularity of cats
as pets has also contributed to large populations of unowned cats [
43
,
48
]. In response
to concerns about their welfare and the problems they may cause (e.g., [
48
,
49
]), since
1990 the Lonely Miaow Association, Inc., Stray Cats New Zealand Trust (hereafter LM;
https://www.lonelymiaow.co.nz (accessed on 1 August 2022)) based in Auckland (New
Zealand’s largest city), has operated a trap–assess–resolve (TAR) approach (also called
‘capture and rehoming’, sensu [
50
]) to manage unowned, free-ranging cats. Volunteers trap
colonies of unowned cats, which are adopted wherever possible or euthanized if they are
unsocialised or in grave ill-health. The aim is to control the population with low euthanasia
without returning cats to life on the streets.
In this paper, we present records from the history of LM’s operations to address:
•Where unowned cat colonies are located.
•Age structures of the trapped cats.
•Outcomes for cats, including analysis by age (kitten or adult).
•Health assessment of cats.
•
Prevalence of neutering amongst unowned cats as a conservative indicator of loss or
abandonment.
•The costs incurred by LM and for what purposes.
We then discuss the advantages and disadvantages of TAR relative to two other
widely-used management options for stray cats: TE and TNR. The data and discussion
form a case study in implementing TAR relevant to urban areas globally where populations
of unowned cats are managed.
2. Materials and Methods
2.1. Context: Managing Unowned Cats in New Zealand
The broad context of animal management in New Zealand, including owned and
unowned cats, is outlined in the New Zealand Animal Welfare Strategy [
51
], (p. 4). The
primary legislation enforcing the strategy is the Animal Welfare Act 1999 [
52
], including
provisions for codes of welfare that, where applied to individual species, state the minimum
standards for care and recommendations for best practices. Sumner and colleagues [
43
]
note that New Zealand legislation is clearer with regard to New Zealand’s feral cats than it
is for stray cats (in New Zealand, as in Australia, feral cats are those remote from human
habitation and with no human interaction [
10
]), so they argue for national legislation for
managing all categories of cats.
2.2. History of Lonely Miaow Association Inc.
LM is a non-profit group that was founded by Peter Dormon in 1990 and incorporated
in 1995. With a general goal of ‘no more strays’, LM runs three main activities across
the Auckland metropolitan area: provision of cat care information, rehoming strays, and
assisting landholders seeking the removal of stray cats.
The rehoming and assistance activities carried out by LM involve TAR in line with
the provisions of New Zealand’s animal welfare code and strategy [
51
,
52
]. Following a
request from the public to remove stray cats, volunteers undertake trapping. If necessary,
they arrange veterinary treatment for trapped cats, including euthanasia for any that
are seriously ill, injured, or too unsocialised for adoption, as judged by both the person
who traps the cats and the veterinarian who examines them (see Supplementary Material
Table S1 for schedules for assessment, plus an example media interview featuring LM
volunteers). Other cats are then placed in foster homes, where ongoing care may include
Animals 2022,12, 2301 4 of 21
socialisation or administering medication/wound dressing under veterinary direction.
All foster carers and their premises are assessed via a home visit and induction before
joining the program. Before being advertised online for adoption, all cats are neutered,
treated for worms and fleas, and given at least their initial vaccinations for panleukopenia
and two strains of cat influenza. Since 2010, following the advent of the New Zealand
Companion Animal Register in 2007, LM began microchipping and registering cats. Today,
all cats are microchipped before adoption and registered in the name of the new owner.
2.3. Collation of Lonely Miaow and Veterinary Records
Summary data tables were extracted by Peter Dormon and Samantha Boston from the
LM database. No data were available for 1990–1995 and only limited data were available
for the period 1995–1998. Assessments focused on:
1.
The site locations where stray cats were trapped (colonies in the LM records) following
requests to remove them by landholders—these were classified as being situated in:
residential properties, commercial premises, countryside/farms, industrial sites, other
(various small categories, including restaurants, hospitals, seaside, council-subsidised
housing, and schools). Data for locations were provided for the date range July 1995
to June 2020.
Other records were provided between July 1999 and December 2019, detailing:
2.
Estimated age—five categories: young kitten (<10 weeks old), older kitten (10 weeks to
6 months), young adult (6 months to 1 year), adult (1–5 years), senior cat
(>5 years old).
3.
Outcome—adopted (including cats adopted by private individuals, adopted by their
fosterers, or taken by pet shops), euthanized, returned to the site, other (including cats
found dead, transferred to other shelters, died during operations or from sickness,
or escaped).
4. Costs—veterinary, food, litter, microchipping, other.
LM uses multiple veterinary clinics across Auckland, with the choice of clinic often
a matter of convenience for LM volunteers. The owner of one of these clinics, the Kohi-
marama Veterinary Clinic, kindly made available the clinical records for LM cats treated in
the clinic in 2000 and between January 2008 and April 2018. We extracted and tabulated
data from these clinical records. All cats not recommended for euthanasia were routinely
wormed, treated for fleas, neutered if they were entire, and given their first vaccinations
for panleukopenia and two strains of cat influenza, so we recorded the prevalence of other
conditions. Data on the health of cats were noted on the date of first presentation, so
that if a cat developed a condition while in foster care that condition was not recorded.
Histories over time could be identified from case numbers or from notes indicating that the
cat had been presented before. We also noted if females were pregnant, lactating, or had
engorged mammaries, as well as any evidence that a cat had been owned previously (e.g.,
already neutered, microchipped). Cats were not routinely checked for feline immunodefi-
ciency virus (FIV) or feline leukaemia virus (FeLV), although when tests were requested,
we recorded the results.
LM also provided receipts for services conducted by a further 26 veterinary clinics
across Auckland between 2010 and 2014 that treated cats at first presentation and during
foster care. The receipts were examined for indications of likely conditions treated by these
clinics, as revealed by the medications used or treatments described.
2.4. Statistical Analyses
Most questions from LM data or veterinary clinical histories were answered with a
crosstabulation of data, including chi-square tests where appropriate. The effects of the
age of the cats (the five age categories), their sex (male or female), and the year in which
they were processed (1999–2019, ignoring the low numbers of cats handled in earlier years)
on the likelihood of adoption were assessed using generalised linear models (GLM) in
SPSS 22 for Windows [
53
]. The probability of adoption was initially modelled using a
Animals 2022,12, 2301 5 of 21
binomial distribution and probit link function, including predictor factors: cat age, cat
sex, year, and age x sex interaction. To further investigate the interaction, a second probit
analysis used an age variable with two levels (kitten and adult) and omitted the sex of the
cats. To explore how the costs of TAR changed with the scale of the operation, Pearson’s
correlation coefficients were calculated relating unit costs to the number of cats processed
annually, with two-tailed tests for differences from zero.
3. Results
3.1. Colonies Resolved, Age Profiles, and Outcomes for Trapped Cats 1995–2020
Between July 1995 and June 2020, 3737 colonies (each defined as a specific callout from
a landholder, usually in response to cats aggregating at a place providing shelter or food)
were processed using TAR. The data cannot determine whether all these colonies were
spatially or temporally independent, so some migration might have been possible between
some of them. The data also include multiple callouts to the same location. Most were on
residential properties (79.1% in 1995–2010, 81.2% in 2011–2020) (Table 1). There was a slight
change in the relative proportions of colonies in the different categories over time, because
of a small drop in the proportion of cats caught at commercial premises or in the peri-urban
environment, and a small increase in the proportion of cats trapped at industrial sites or
council-subsidised housing in 2011–2020 (χ26= 54.9, p< 0.001).
Table 1.
Cat colonies (total n = 3737) resolved by Lonely Miaow, Auckland, New Zealand, between
June 1995 and June 2020.
Year Range: July 1995–December 2010 January 2011–June 2020
Location of Colony Number (Percent) of Colonies Number (Percent) of Colonies
Residential property 1300 (79.1%) 1700 (81.2%)
Commercial premises 164 (10.0%) 154 (7.4%)
Peri-urban 69 (4.2%) 44 (2.1%)
Industrial site 29 (1.8%) 52 (2.5%)
Council subsidized housing 10 (0.6%) 57 (2.7%)
Hospital 15 (0.9%) 5 (0.2%)
Other (e.g., schools, beach, restaurants) 56 (3.4%) 82 (3.9%)
Sub-total 1643 2094
In total, 14,611 cats were processed through LM between July 1995 and December 2019.
The mean number of cats processed per annum was 695
±
332 s.d. (calculated from 1999
when annualised records began). The overall sex ratio was 45.6% males and 54.4% females
for the 13,265 cases where sex was recorded. The sex ratio did not change significantly over
time (
χ221
= 16.10, p= 0.762). However, the sex ratio did change with the age of the animal.
There was an equal sex ratio for young kittens (50.0% male). This shifted to a bias towards
females for young adults (32.5% male) and adults (37.8% male), before returning to a bias
to males for seniors (58.9% male) (χ24= 227.16, p< 0.001).
Age data were available for most (98.8%) cats. Overall, c. two-thirds were kit-
tens: 51.9% as young kittens (<10 weeks) or 16.1% as older kittens (10 weeks–6 months).
The remaining third were adults: 9.2% young adults (6 months–1 year), 19.7% adults
(1 year–5 years), and 1.9% seniors (>5 years).
There were 6 years when the number of cats processed annually reached or ex-
ceeded ~1000: 2000, 2007, 2008, and 2015–2017 (Figure 2a). During these years, there
were marginally more adult cats (34.1%) processed through the system than the overall
value (30.8%), while kittens averaged 64.2% compared to the overall value of 68.0% of all
cats processed (Figure 2b).
Animals 2022,12, 2301 6 of 21
Figure 2.
The age of cats processed for trap–assess–resolve (TAR) through Lonely Miaow, Auckland,
New Zealand, between July 1995 and December 2019, showing the breakdown by year. Rectangles
at the top highlight 6 years where efforts exceeded ~1000 cats per annum. The exact numbers are
shown in (a) and the percentages in (b).
The fate of cats processed through LM shifted markedly over time, with the proportion
of cats being adopted nearly doubling from 43.4% of cats in the first decade of establishment
to 72.5% over the last five years (Figure 3). During this time, the proportion of cats
euthanized decreased from 40.0% in the first decade of establishment to only 12.1% over
the last five years.
Animals 2022,12, 2301 7 of 21
Figure 3.
The fate of cats processed for trap–assess–resolve (TAR) through Lonely Miaow, Auckland,
New Zealand, between July 1995 and December 2019, showing the breakdown by year. ‘Adopted’
includes adoptions brokered by Lonely Miaow, cats adopted by a foster carer, or cats placed with a
pet shop. ‘Other ’ includes cats that died, were found dead, escaped, or were transferred to another
shelter, and ‘unknown’ where a volunteer had not entered the data. The exact numbers are shown
in (a) and the percentages in (b).
Younger cats were more likely to be adopted than older cats (probit GLM model
including cat age, cat sex, year, and age x sex interaction: Wald chi-square df 4 = 2285.2,
p< 0.001), and annual adoptions increased over time (Wald chi-square df 1 = 179.3,
p< 0.001). Overall, females were more likely to be adopted (Wald chi-square df
1 = 6.9,
Animals 2022,12, 2301 8 of 21
p= 0.009). The adoption rates of female cats versus male cats changed with age, with male
kittens more likely to be adopted than female kittens and adult females more likely to be
adopted than adult males (Wald chi-square df 4 = 30.0, p< 0.001). To further investigate
the interaction, a second probit analysis used an age variable with two levels (kitten and
adult) and treated time as a categorical variable with four periods each of five years. We
omitted the sex of the cats. It confirmed strongly that kittens were more likely to be adopted
than adults (Wald chi-square df 1 = 1758.9, p< 0.001), annual adoptions changed with the
period (Wald chi-square df 3 = 525.2, p< 0.001), and the adoption rate of kittens increased
more rapidly with time than adults (Wald chi-square df 3 = 50.3, p< 0.001) Figure 4. Ex-
amining kittens and adults separately, the effect of sex was significant in kittens, where
82.0% of females were adopted as opposed to 84.1% of males (Wald’s chi-square = 8.5, df 1,
p= 0.003). In adults, sex was also significant (Wald’s chi-square = 13.2, df = 1, p= 0.001),
with 42.5% of females adopted, as opposed to 36.7% of males, a reversal of the previous bias.
Figure 4.
The proportion of kittens and adult cats adopted between 1999 and 2019 through the
trap–assess–resolve (TAR) program run by Lonely Miaow, Auckland, New Zealand. Open symbols
represent kittens and filled symbols represent adults. Error bars represent 95% confidence intervals.
3.2. Health Assessments
Excluding microchipping, neutering, flea and worm treatments, and vaccinations,
541 (21.9%) of LM cats who presented to the Kohimarama Veterinary Clinic in 2000 and
2008–2020 required veterinary treatment on the first visit (Figure 5). Common ailments
included symptoms of respiratory tract infections indicative of cat influenza (eye or nasal
discharges, rattly chest, sneezes) (61.7%), trauma including fractures, amputations, gloving
injuries (where the skin is stripped from a limb or the tail) and fight wounds (14.4%), dental
problems, including gingivitis, broken teeth, and periodontitis (9.8%), and ringworm (9.2%).
Overall, 63 cats were tested for FIV (41.3% tested positive) and 17 for FeLV (23.5% tested
positive). Given that not every cat was tested, these results do not indicate the prevalence of
FIV and FeLV in the Auckland stray cat population. A total of 80 (15.0%) of the 452 female
cats were neutered although pregnant, aborting the kittens. A further 37 females (8.2%) had
Animals 2022,12, 2301 9 of 21
recently given birth because they had engorged mammaries indicative of recent feeding of
kittens. Every effort was made to trap kittens with their mothers.
Figure 5.
Ailments for which 541 of 2470 cats processed for trap–assess–resolve (TAR) were treated
by the Kohimarama Veterinary Clinic in 2000 and between 2008 and 2018 as part of the processing for
Lonely Miaow, Auckland, New Zealand. Note that a cat could be treated for multiple conditions,
so the sum of the bars in the figure exceeds 541.
Receipts from a further 26 veterinary practices between 2010 and 2014 showed that
the main conditions for cats at first presentation or in foster care were:
•Respiratory and eye infections indicative of cat influenza.
•Skin infections, mainly ringworm.
•Urinary tract infections.
•
Severe worm infestations requiring repeated and intensive worming, often associated
with diarrhoea and dehydration.
•Secondary bacterial infections associated with cat influenza or diarrhoea.
There was evidence of significant discounting of fees on most occasions.
3.3. Estimation of Abandonment Rates
A total of 31 cats (1.2% of initial presentations to the Kohimarama Veterinary Clinic in
2000 and 2008–2018) were identified as having been owned previously, because when they
were trapped they were already neutered and were not ear-tipped or tattooed to identify
them as part of a TNR program (26 cats), had a microchip (2 cats), had dental work that had
evidently been done by a veterinarian (1 cat), was judged so friendly and socialised that it
must have been owned recently (1 cat), or the owner was traced (method not clear, 1 cat).
Given that these cats must have been lost or abandoned, they gave a minimum baseline
indication of loss or abandonment in the Auckland stray cat population. While this figure
is low, the real rate will be increased by the loss or abandonment of entire animals that are
not microchipped, which could not be detected.
Animals 2022,12, 2301 10 of 21
3.4. Costs
Between 1996 and 2017, NZD 2.1 million was spent supporting the TAR program.
Annual expenditure grew from NZD 8879 in 1996 to NZD 269,451 in 2017. Most costs
(61.1%) were veterinary (Figure 6). However, we note that the contributions of unpaid
volunteers, donations of consumables, such as cat litter, and the pro bono or discounted
work of some veterinarians are not included in these figures. Estimates of volunteer hours
and pro bono veterinary contributions could be assessed by asking participants to keep
diaries of hours, which could then be costed at an agreed rate.
Figure 6.
The financial costs of processing 13,062 stray cats (where reliable data were available)
processed for trap–assess–resolve (TAR) through Lonely Miaow, Auckland, New Zealand, between
1996 and 2017, (a) by category of cost and (b) average estimated per cat.
From 1999 (when annualised case records began) to 2017, the average costs per cat rose
from NZD 58 to NZD 234 (Figure 6inset). Food and litter costs per cat did not change with
the annual number of cats processed (Figure 7a, r = 0.049, p= 0.853). Veterinary costs per
cat increased strongly (Figure 6a), suggesting that more thorough care was being provided,
the prevalence or severity of ailments rose, veterinary costs rose at around triple the general
rate of inflation (RBNZ 2021), or that there was reduced fee discounting or pro bono work.
Total and veterinary costs per cat were not related to the number of cats processed each
year (Figure 7b,c total: r = 0.113, p= 0.646 vet: r = 0.109, p= 0.657). This indicates that as
LM grew, there were, in net, no substantial economies of scale; either positive (e.g., bulk
discounts) or negative (e.g., saturating the capacity of pro bono services).
Animals 2022,12, 2301 11 of 21
Figure 7.
Per capita costs in relation to the number of cats processed for trap–assess–resolve (TAR)
through Lonely Miaow, Auckland, New Zealand. Each point represents a year, from 1999–2017.
(
a
) gives food and litter costs, (
b
) gives veterinary costs and (
c
) gives total costs. All costs are in NZD.
Animals 2022,12, 2301 12 of 21
4. Discussion
The data arising from the LM program are relevant in terms of assessing the welfare
of stray cats in Auckland, as well as evaluating the advantages and disadvantages of TAR
relative to TNR and TE in terms of costs and the range of problems related to stray cats
that they solve. These data represent detailed insights into the lives of stray cats on the
streets of a developed country recognised for its high value in animal welfare [
51
,
52
]. The
study also captures the enormous efforts of veterinary professionals, volunteers, providers
of pro bono services, and an enduring not-for-profit organisation in neutering, medicating,
and rehoming stray cats.
4.1. Distribution of Stray Cats in Auckland
LM responds to requests from the public to remove stray cats, so the distribution of
colonies likely reflects, to an unknown extent, the distribution of complaints to areas of a
higher population rather than the distribution of cats. Most LM colonies over the period
1995–2020 came from residential areas (81% for 2011–2020 and 79% for 1995–2010 in the
present study; 75% for 1991–2011 [
54
]. Thus, there is either a concentration of colonies in
residential areas over time or increased reports of colonies there relative to other areas. The
increase in the number of colonies reported over time could reflect increased awareness of
LM services, increased public awareness of stray cats and concern for them, or possibly
increases in stray cat numbers.
Separate cat collection records in Auckland revealed that strays were geographically
clustered, with possibly a higher prevalence in economically-deprived areas [
54
]. In that
study, colonies were associated with residential areas of higher human population density
and, more weakly, higher levels of social deprivation that could be targeted with education
campaigns regarding the management of stray cats, as well as subsidies to assist in trapping,
neutering, and rehoming. Particular attention could be given to any problems caused by
provisioning these cats. The idea of identifying communities where education would be
effective could be applied internationally, with a growing body of education approaches
addressing barriers to good husbandry (e.g., [55–59]).
4.2. Welfare of Stray Cats in Auckland
The welfare of stray cats is an important concern in deciding whether and how to
manage their populations. Visual assessments of cats conducted at varying distances
are commonly used to note such factors as body conditions, coat conditions, injuries, and
prominent eye or nasal discharges (e.g., [
6
,
60
]). However, they are less commonly calibrated
against actual veterinary examinations as proposed by [
48
]. Where detailed records are
kept of the health of cats admitted to shelters or processed in TNR programs [
61
,
62
],
or cadavers are examined following euthanasia (e.g., [
7
]), data indicate compromised
animal welfare. Therefore, although some authors concluded that the managed stray cats
were in good health (e.g., [
63
]), other studies reveal concerningly high percentages of cats
in poor condition (e.g., [64,65]).
Zito et al. [
48
] used distance observation to assess the condition of a convenience
sample of 676 cats (divided into n = 213 companion cats, n = 210 managed stray cats,
and n = 253
unmanaged stray cats) from ‘various unspecified locations’ across Auckland.
They identified the health conditions for managed and unmanaged stray cats: eye and
nasal discharges, and injuries (‘trauma’ in the present study). Crusting around the ears
was reported in 11.4% of stray cats [
48
] but was not one of the major categories of health
problems in LM cats (present study). In contrast, snuffles, rattly chests, ringworm infections,
and dental problems were commonly reported in the LM cats, as well as some cats that
were FIV- or FeLV-positive, but these conditions were not recorded by [
48
]. Clinical notes
for the LM cats indicated the diversity and severity of less common conditions, for example:
“Covered in fleas, more fleas than cat. Smaller (cat) dehydrated and both anaemic”. Records also
sometimes included comments on the fate of cats that did not reach the clinic, for exam-
ple: “post-partem (sic) cat spey, left flank as mammary tissue engorged (kittens killed by a dog)”.
Animals 2022,12, 2301 13 of 21
Zito et al. [
66
] also reported that 23 of 429 cats (5.4%) trapped in their TNR trial were
euthanized for one or more conditions, including ataxia, diarrhoea, oral ulcers, ringworm,
being underage, upper respiratory tract infections, trauma, cancer, corneal ulcers, dehydra-
tion, emaciation, or FIV. Although the prevalence of euthanasia in [
66
] was less than that
reported for LM, overall, the greater range of conditions requiring intervention when cats
were handled cautions against using visual assessments alone.
The consistent pattern over 20 years of c. 80% of cats being less than a year old,
and only c. 2.0% estimated to be over 5 years old, indicates short life expectancies or
possibly trap-shyness in older cats. The high removal of older cats is less likely to be
a factor, given that our data indicate that kittens are more likely to be rehomed. This
low survival rate agrees with estimates of 75–90% mortality by six months of age for
free-ranging kittens from high-density populations [
7
,
62
,
67
–
69
]. Sex ratios were close
to parity in young kittens, but by adulthood were biased to females, before reverting to
parity or male-biased ratios in older cats. This is consistent with roaming and fighting in
males leading to higher mortality at early ages (e.g., data on the male-biased prevalence
of FIV in stray or shelter cats, [
20
,
61
,
70
,
71
]), while older females might be exhausted by
constant breeding (c. 23% of female LM cats neutered at the Kohimarama Veterinary Clinic
were either pregnant or showed signs of supporting kittens). The maintenance of kitten
numbers—despite TAR—implies migration, ongoing abandonment of cats, small numbers
of cats not removed, breeding successfully, or that the number of cats processed was small
relative to the population.
By contrast, the age profiles of owned cats include larger proportions of cats over five
years old (e.g., [
72
–
74
]). However, even despite the care of their owners, pet cats display
higher risk behaviours when young and are more likely to suffer trauma (Auckland: [
75
],
elsewhere: [
17
]). This supports our interpretation of high mortality of young stray cats.
Alternative explanations are that calls for LM to help are more likely in response to sightings
of kittens; however, this would not explain the age-related shifts in the sex ratio of trapped
cats. An alternative is that older animals are wary and trap-shy and, hence, less likely to be
caught; this hypothesis requires further data on trappability.
Based on the LM data, we argue that there is a strong case on welfare grounds to
reduce Auckland’s stray cat population. 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].
4.3. Comparing TAR, TNR, and TE
Given the significant welfare issues for stray cats themselves and the nuisance, public
health risks and depredation of wildlife they may cause, intervention is needed. Modelling
studies indicate that, regardless of the choice of management strategy, the intensity of
application is critical to achieving a strong population reduction [
44
,
79
]. In this context,
key features of TE, TNR, and TAR are summarised in Table 2.
If speedy population reduction is desired, then TE and TAR have the greatest potential
for rapid results because cats are removed immediately. While adoption is an important
component of TNR, unadopted cats are still returned, so a significant population reduction
may be delayed or unachievable. Whether this is acceptable will depend on the urgency
of resolving problems, such as wildlife impacts or public health risks, which are not
mitigated by neutering alone [
25
,
80
]. TNR is attractive to many because euthanasia is
reduced [
8
,
43
], although the prevalence of desexing pregnant cats may be higher than
many realise [
81
]. This is an ethical concern [
82
], alongside the welfare of returned cats
irrespective of whether or not there is caregiver support [
77
,
78
,
83
]. Whether TAR reduces
public or veterinary health risks or reduces predation by cats, depends on the husbandry of
the adopted cats by their new owners. If the cats are allowed to roam unrestricted, benefits
Animals 2022,12, 2301 14 of 21
may be limited [
24
,
29
]. Finally, adoption is assumed to be for the rest of the cat’s life [
84
].
Post-adoption follow-up studies could address this question for adoptions under both
TAR and TNR (e.g., [
85
,
86
]), including the possibility that cats are assessed incorrectly as
suitable for adoption.
Comparative data on the costs of different approaches to stray cat management are
limited and complicated by whether or not they include costs for: ongoing maintenance
of TNR colonies over the lifespans of the returned cats, discounting or pro bono work
by veterinarians, volunteer labour, community education to increase desexing of owned
pets and reduce abandonment, and valuing the benefits of particular interventions against
costs [
37
,
38
,
87
]. Valuations of benefits are contentious, especially when values are placed
on the wildlife lost to predation or cat-borne diseases [
88
]. Nevertheless, potential benefits
that might be assessed are saved costs to public health, veterinary health, agricultural
production from the curtailment of diseases transmitted by stray cats [
89
], wildlife conser-
vation [
25
–
27
], and the improved mental health of animal welfare professionals who can be
traumatised by frequently euthanising healthy animals [
90
,
91
]. Overall, the costs reported
in the literature at best provide comparisons for a local situation. Before generalising,
agreement is necessary on the costs of subsidies and volunteer efforts (including ongoing
maintenance where provisioning of TNR colonies is established), benefit valuations, and
the projected life of a control program.
Table 2.
Comparisons of the main features of trap and euthanize (TE), trap–neuter–return (TNR),
and trap–assess–resolve (TAR) for managing populations of unowned cats.
Feature TE TNR TAR
Speed of population
reduction
Rapid extirpation in
closed populations but
needs reapplication if
abandonment or
migration replenishes
cats [37,87].
Some local successes are claimed,
especially where populations are closed
and there is high adoption within the
program (e.g., [92,93]); however,
Gunther et al. [60] observed that
achieving sterilization rates of at least 75%
(required for population decline) is ‘almost
impossible to reach and sustain on a
meta-population scale.’
Gunther et al. [46]
reported population reductions of
approximately 7% per year following
high intensity (>70% neutering)
maintained across contiguous sites
covering a 20 km2urban area.
Should be rapid in a closed
population because cats are
removed. Needs reapplication
when numbers are replenished
by migration or abandonment,
similar to the problems noted
for TE and for TNR that are not
applied across contiguous areas
simultaneously [46].
Addresses problems
caused by stray cats
Yes, if applied at a level
that reduces populations.
Euthanized cats cannot
breed, experience poor
welfare, threaten wildlife,
cause nuisance, or
spread disease.
From a welfare perspective, prevents the
birth of kittens likely to die young [44],
but high numbers of kittens may be
aborted when neutering pregnant
queens [81]. May reduce disease
transmission if it includes vaccination [
6
].
Neutered cats are less likely to fight or
cause nuisances [6,63]. They still hunt,
though, so that problem is
unaddressed [25,26,46,94]. Some cats are
adopted, gaining lasting care.
Provides veterinary care
including vaccination and, for
adoptable cats, provides lasting
care in homes. The husbandry
of the adopted cats will
determine the levels of
nuisances they may cause, the
risk of them spreading diseases,
and any threat to wildlife. Seo
et al. [83,95] recommended
adoption as a replacement for
TNR for animal welfare and
public health reasons.
Opportunities for
citizens to be
involved
None, assuming that
citizens are unlikely to
volunteer for trapping
followed by euthanasia.
High, including trapping and
transporting cats and providing food and
shelter to colonies.
High, including trapping and
transporting cats and fostering.
Animals 2022,12, 2301 15 of 21
Table 2. Cont.
Feature TE TNR TAR
Euthanasia
All animals (other than
pets returned to owners)
are euthanized.
Greatly reduced. Only ill cats
are euthanized.
Greatly reduced. Only ill or
unsocialized cats are
euthanized. However, there is a
risk that timid cats are
euthanized unnecessarily or
that unsuitable cats are adopted.
Other ethical
considerations
Fits within a utilitarian or
consequentialist ethical
approach, in which
managers strive to achieve
the best outcomes overall
for all animals
involved [96,97], which in
this case would include
the cats themselves, other
organisms or people
threatened with diseases,
and wildlife at risk
of predation.
Follows a deontological or rights
approach respecting intrinsic animal
rights, although ethically TNR must also
justify neutering [
98
] and death of kittens
when neutering pregnant queens [82].
TNR may also encourage dumping cats
at TNR colonies [99,100]. Even when fed,
cats returned to the site may have
compromised welfare [7], see also this
paper. TNR also values cats as a species
over the wildlife they may hunt or infect
with disease [101].
Similar to TNR in following a
deontological approach, as well
as needing to justify neutering
(including of pregnant animals)
from an ethical perspective [
82
,
98
].
Adopting cats, rather than
returning them to the site,
should provide a high level of
care. Euthanasia of unadoptable
cats can be argued to be
preferable to the risks associated
with returning them [102].
Whether adopted cats still
threaten wildlife or spread
disease is dependent on their
husbandry.
Moral distress to
veterinary and
animal welfare
professionals
Moral distress caused by
euthanasia or leaving stray
cats on the streets are only
addressed if populations
are suppressed long-term.
Euthanasia is greatly reduced but distress
over unowned cats remains. Some moral
stress may arise from neutering pregnant
queens [82].
Euthanasia is greatly reduced
(but not as much as TNR), while
any distress over returning cats
to the streets is eliminated.
Some moral stress may arise
from neutering pregnant
queens [82].
Costs per cat
USD 52–123 [38].
USD 215.82 † [37].
~Half the cost of TNR
according to comparative
modelling [87].
USD 20–97 [38].
~Twice the cost of TE according to
comparative modelling [87].
At least USD 45 [46], based on a cost of
over USD 1 million for a program
sustained over 9 years in a 20 km2urban
area that neutered 22,144 cats.
USD 104–550 [38].
USD 103.98 ‡ (this paper).
† AUD 277.5 at exchange rate (16 May 2021); ‡ NZD 142.50 at exchange rate (26 May 2021).
TAR is not without potential problems. Timid cats may be euthanized unnecessarily
instead of proposed for adoption, or some cats may be recommended for adoption when
they are temperamentally unsuited to be pets. The extent to which TAR ameliorates public
health risks or hunting behaviour also depends on the husbandry of adopted cats. If they
are kept on the owners’ properties, public health is enhanced, and hunting is curtailed.
If adopted cats are allowed to roam freely, they may still pose health risks, hunt wildlife,
and risk physical trauma themselves.
4.4. Integrated Responses to Reduce Populations of Stray Cats
Despite removing over 14,000 stray cats from the streets of Auckland, there is no sign
of a decline in the need for LM services. We have no data showing reduced numbers of cats
at managed locations, reduced numbers of locations with cats, or declining captures per
unit effort. Thus, programs such as TAR should be implemented as part of a wider strategy,
ideally conducted at the level of the municipality and involving all stakeholders, addressing
the management of owned animals to prevent unwanted breeding, abandonment, and loss
while simultaneously reducing the number of strays. One example of such an integrated
approach is the Australian Capital Territory’s 2021–2031 Cat Plan [
103
], which seeks to
achieve the vision that by 2031 ‘All cats in the ACT will be owned, wanted and cared for by
Animals 2022,12, 2301 16 of 21
responsible owners.’ Action 8 under that plan requires: ‘Work with animal care and rescue
organisations to manage semi-owned and unowned cats in public places, through trap,
de-sex and adopt activities’—TAR, as described in this paper.
The primary need is to reduce unwanted breeding. Neutering of owned animals,
including increased use of early-age or prepubertal neutering (before six months) and
mandatory neutering prior to the sale or transfer of registered animals, will prevent un-
planned breeding and possible abandonment of unwanted animals [
104
,
105
]. Many of
the concerns regarding the effects of neutering on the health and development of cats
were rebutted with empirical data in the publications arising from Belgium’s Sterycat Pro-
gram [
106
–
109
]; however, there remains evidence that even in countries reporting a high
prevalence of neutering, many cats are not neutered until later in life [
110
–
112
]. Groups
such as Australia’s National Neutering Network (https://ndn.org.au) or the United King-
dom’s Kitten Neutering Database (KiND, http://www.kind.cats.org.uk (31 August 2022))
may help to reverse this trend. Sumner et al. [
43
] report that the New Zealand Veterinary
Association endorses prepubertal desexing.
Mandatory IDs, including both microchipping and collar-worn IDs, will assist by
improving the low rates of returning lost cats to their owners [
113
,
114
]. Measures can
also be taken to encourage people to own and keep pets, with signs that the adoption
of a shelter cat is becoming desirable, at least in some countries [
115
]. There was even
a surge in pet adoptions in relation to the COVID-19 pandemic [
116
]. More pet-friendly
rental arrangements, better provision of information and services to owners (for example,
regarding the welfare benefits of containment of cats on their owners’ property), as well as
assistance to shelters in rehoming stray animals, may also assist [
55
,
57
,
84
]. If people are
confident that cats surrendered to shelters are highly likely to be rehomed, they may not
feed strays and thereby encourage unwanted breeding. Thus, rather than seek positive
attitudes towards unowned cats alone [
117
], we would prefer that citizens have positive
attitudes towards finding homes for unowned cats. We are divided as to whether it is ever
good practice to feed strays. Some of us disagree with feeding because it may encourage the
abandonment of animals at feeding stations, facilitate recruitment, and encourage rodents
and other pests that take scraps. Others argue that feeding alleviates hunger and, although
it makes trapping more difficult, it is better than cats suffering from starvation. LM neither
encourages nor discourages people to feed strays.
In our opinion, TAR fits well within such an integrated strategy. In comparison to TE,
it reduces euthanasia and addresses the concern expressed by some citizens regarding lethal
control, although we acknowledge that we have no evidence for population reduction.
Nevertheless, in comparison to TNR, it rehomes stray cats, potentially reducing a wide
range of welfare problems and, with appropriate husbandry, concerns over nuisances,
public health, and attacks on wildlife. Ultimately, the choice of approach in any situation
rests with the local community [83,100].
5. Conclusions
The poor animal welfare outcomes, public health risks, and threats to wildlife often
associated with unowned, free-roaming cats, demand action [
8
,
36
]. While this can be
presented as a choice between euthanasia and TNR, the work of LM shows that TAR can
also be considered. TAR reduces euthanasia considerably relative to culling, and adoption
should raise the level of care well above life on the street. TAR’s effectiveness would
be enhanced by education on the value of pre-pubertal desexing [
118
] and the problems
caused by abandoning cats [
83
,
100
]. Further data are needed on its success in reducing
populations and curtailing risks to wildlife and public health.
Supplementary Materials:
The following supporting information can be downloaded at: https://
www.mdpi.com/article/10.3390/ani12172301/s1, Table S1: Examples of forms used over the years to
assess the health and temperament of trapped cats.
Animals 2022,12, 2301 17 of 21
Author Contributions:
Conceptualization, M.C.C., P.D., P.A.F.; formal analysis, M.C.C., J.S.B., P.A.F.,
F.R.S.; resources, P.D., S.B.; writing—original draft preparation, M.C.C., P.D., P.A.F., J.S.B., S.B., H.M.C.,
F.R.S., all authors; writing—review and editing, all authors. All authors have read and agreed to the
published version of the manuscript.
Funding: This research received no external funding.
Institutional Review Board Statement:
Ethical review and approval were not required for the
animal component of this study because no animals were handled specifically for the purpose of this
study. Murdoch University’s Human Research Ethics Committee exempted this project from needing
Human Ethics approval on 6 May 2021, approval number HREC 2021/066.
Informed Consent Statement: Not applicable.
Data Availability Statement:
Most relevant data are reproduced in the figures and tables within the
paper. The original Lonely Miaow (Inc.) data are confidential, as are the original case records we
were permitted to examine for the study.
Acknowledgments:
We thank the volunteers at Lonely Miaow, Inc. for their patience in downloading
data and reports as well as answering our many questions. Russell Tucker, owner of the Kohimarama
Veterinary Clinic, generously made available all invoices and clinical histories related to Lonely
Miaow, Inc.’s work between 2008 and 2018 to enable the assessment of the health of the rescued cats.
John Read kindly commented on an earlier version of the paper, although we do not imply that he
agrees with all our opinions. Belinda Cale drew Figures 2–4and 7. The manuscript was enhanced by
the perceptive comments of two reviewers.
Conflicts of Interest:
Peter Dormon and Samantha Boston are volunteers at Lonely Miaow (Inc.).
The other authors declare no conflict of interest.
References
1. Spotte, S. Free-Ranging Cats: Behavior, Ecology, Management; John Wiley and Sons: Chichester, UK, 2014.
2.
Fraser, G.; Huang, Y.; Robinson, K.; Wilson, M.S.; Bulbulia, J.; Sibley, C.G. New Zealand pet owners’ demographic characteristics,
personality, and health and wellbeing: More than just a fluff piece. Anthrozoos 2020,33, 561–578. [CrossRef]
3.
Krittanawong, C.; Kumar, A.; Wang, Z.; Jneid, H.; Virani, S.S.; Levine, G.N. Pet ownership and cardiovascular health in the US
general population. Am. J. Cardiol. 2020,125, 1158–1161. [CrossRef] [PubMed]
4.
McKay, S.A.; Farnworth, M.J.; Waran, N.K. Current attitudes toward, and incidence of, sterilization of cats and dogs by caregivers
(owners) in Auckland, New Zealand. J. Appl. Anim. Welf. Sci. 2009,12, 331–344. [CrossRef] [PubMed]
5.
Peterson, M.N.; Hartis, B.; Rodriguez, S.; Green, M.; Lepczyk, C.A. Opinions from the front lines of cat colony management
conflict. PLoS ONE 2012,7, e44616. [CrossRef]
6.
Gunther, I.; Raz, T.; Klement, E. Association of neutering with health and welfare of urban free-roaming cat population in Israel,
during 2012-2014. Prev. Vet. Med. 2018,157, 26–33. [CrossRef]
7.
Crawford, H.M.; Calver, M.C.; Fleming, P.A. Subsidised by junk foods: Factors influencing body condition in stray cats (Felis catus).
J. Urban Ecol. 2020,6, juaa004. [CrossRef]
8.
Wolf, P.J.; Hamilton, F. Managing free-roaming cats in U.S. cities: An object lesson in public policy and citizen action. J. Urban Aff.
2020,44, 221–242. [CrossRef]
9.
Zito, S.; Morton, J.; Paterson, M.; Vankan, D.; Bennett, P.C.; Rand, J.; Phillips, C.J.C. Cross-sectional study of characteristics of
owners and nonowners surrendering cats to four Australian animal shelters. J. Appl. Anim. Welf. Sci.
2016
,19, 126–143. [CrossRef]
10.
Lepczyk, C.A.; Calver, M.C. Cat got your tongue? The misnomer of ‘community cats’ and its relevance to conservation. Biol.
Invasions 2022,24, 2313–2321. [CrossRef]
11.
Hamilton, F. Implementing Nonlethal solutions for free-roaming cat management in a county in the southeastern United States.
Front. Vet. Sci. 2019,6, 259. [CrossRef]
12.
Legge, S.; Murphy, B.P.; McGregor, H.; Woinarski, J.C.Z.; Augusteyn, J.; Ballard, G.; Baseler, M.; Buckmaster, T.; Dickman, C.R.;
Doherty, T.; et al. Enumerating a continental-scale threat: How many feral cats are in Australia? Biol. Conserv.
2017
,206, 293–303.
[CrossRef]
13.
Farnworth, M.J.; Muellner, P.; Benschop, J. A Systematic Review of the Impacts of Feral, Stray and Companion Domestic Cats (Felis catus)
on Wildlife in New Zealand and Options for Their Management; Unitec Institute of Technology: Auckland, New Zealand, 2013.
14.
Rochlitz, I. Clinical study of cats injured and killed in road traffic accidents in Cambridgeshire. J. Small Anim. Pract.
2004
,45,
390–394. [CrossRef]
15.
Rochlitz, I. Feline welfare issues. In The Domestic Cat: The Biology of its Behaviour; Turner, D.C., Bateson, P., Eds.; Cambridge
University Press: Cambridge UK, 2014; pp. 131–154.
Animals 2022,12, 2301 18 of 21
16.
Calver, M.C.; Adams, G.; Clark, W.; Pollock, K.H. Assessing the safety of collars used to attach predation deterrent devices and ID
tags to pet cats. Anim. Welf. 2013,22, 95–105. [CrossRef]
17.
McDonald, J.L.; Cleasby, I.R.; Brodbelt, D.C.; Church, D.B.; O’Neill, D.G. Mortality due to trauma in cats attending veterinary
practices in central and south-east England. J. Small Anim. Pract. 2017,58, 570–576. [CrossRef] [PubMed]
18.
Moreau, D.; Cathelain, P.; Lacheretz, A. Comparative study of causes of death and life expectancy in carnivorous pets (II). Rev.
Med. Vet. 2003,154, 127–132.
19.
Gehrt, S.D.; Wilson, E.C.; Brown, J.L.; Anchor, C. Population ecology of free-roaming cats and interference competition by coyotes
in urban parks. PLoS ONE 2013,8, e75718. [CrossRef]
20.
Hosie, M.J.; Addie, D.; Belák, S.; Boucraut-Baralon, C.; Egberink, H.; Frymus, T.; Gruffydd-Jones, T.; Hartmann, K.; Lloret, A.;
Lutz, H.; et al. Feline immunodeficiency ABCD guidelines on prevention and management. JFMS
2009
,11, 575–584. [CrossRef]
21.
Nation, P.N.; St. Clair, C.C. A forensic pathology investigation of dismembered domestic cats: Coyotes or cults? Vet. Pathol.
2019
,
56, 444–451. [CrossRef]
22.
Bonnington, C.; Gaston, K.J.; Evans, K.L. Fearing the feline: Domestic cats reduce avian fecundity through trait-mediated indirect
effects that increase nest predation by other species. J. Appl. Ecol. 2013,50, 15–24. [CrossRef]
23.
Mori, E.; Menchetti, M.; Camporesi, A.; Cavigioli, L.; Tabarelli de Fatis, K.; Girardello, M. License to kill? Domestic cats affect a
wide range of native fauna in a highly biodiverse Mediterranean country. Front. Ecol. Evol. 2019,7, 477. [CrossRef]
24.
Legge, S.; Woinarski, J.; Dickman, C.; Murphy, B.P.; Woolley, L.-A.; Calver, M.C. We need to worry about Bella and Charlie: The
impacts of pet cats on Australian wildlife. Wildl. Res. 2020,47, 523–539. [CrossRef]
25.
Greenwell, C.N.; Calver, M.C.; Loneragan, N.R. Cat gets its tern: A case study of predation on a threatened coastal seabird.
Animals 2019,9, 445. [CrossRef] [PubMed]
26.
Nagata, J.; Haga, A.; Kusachi, Y.; Tokuyoshi, M.; Endo, H.; Watari, Y. Cats were responsible for the headless carcasses of
shearwaters: Evidence from genetic predator identification. Mammal Study 2022,47, 197–204. [CrossRef]
27.
Kristancic, A.R.; Kuehs, J.; Beal Richardson, B.; Baudains, C.; Hardy, G.E.S.; Fleming, P.A. Biodiversity conservation in urban
gardens—Pets and garden design influence activity of a vulnerable digging mammal. Landsc. Urban Plan.
2022
,225, 104464.
[CrossRef]
28.
Lepczyk, C.A.; Lohr, C.A.; Duffy, D.C. A review of cat behavior in relation to disease risk and management options. Appl. Anim.
Behav. Sci 2015,173, 29–39. [CrossRef]
29.
Must, K.; Lassen, B.; Jokelainen, P. Seroprevalence of and risk factors for Toxoplasma gondii infection in cats in Estonia. Vector-Borne
Zoonotic Dis. 2015,15, 597–601. [CrossRef] [PubMed]
30.
Brosh-Nissimov, T.; Ben-Ami, R.; Astman, N.; Malin, A.; Baruch, Y.; Galor, I. An Outbreak of Microsporum canis infection at a
military base associated with stray cat exposure and person-to-person transmission. Mycoses 2018,61, 472–476. [CrossRef]
31.
Halánová, M.; Petrová, L.; Halán, M.; Trbolová, A.; Babinská, I.; Weissová, T. Impact of way of life and environment on the
prevalence of Chlamydia felis in cats as potentional sources of infection for humans. AAEM 2019,26, 222–226. [CrossRef]
32.
Chalkowski, K.; Wilson, A.E.; Lepczyk, C.A.; Zohdy, S. Who let the cats out? A global meta-analysis on risk of parasitic infection
in indoor versus outdoor domestic cats (Felis catus). Biol. Lett. 2019,15, 20180840. [CrossRef]
33.
Sandøe, P.; Nørspang, A.P.; Kondrup, S.V.; Bjørnvad, C.R.; Forkman, B.; Lund, T.B. Roaming companion cats as potential causes
of conflict and controversy: A representative questionnaire study of the Danish public. Anthrozoos
2018
,31, 459–473. [CrossRef]
34. Robertson, S.A. A review of feral cat control. JFMS 2008,10, 366–375. [CrossRef] [PubMed]
35.
Crawford, H.M.; Calver, M.C.; Fleming, P.A. A case of letting the cat out of the bag—why trap-neuter-return is not an ethical
solution for stray cat (Felis catus) management. Animals 2019,9, 171. [CrossRef]
36.
Calver, M.C.; Crawford, H.M.; Fleming, P.A. Response to Wolf et al.: Furthering debate over the suitability of trap-neuter-return
for stray cat management. Animals 2020,10, 362. [CrossRef] [PubMed]
37.
Read, J.L.; Dickman, C.R.; Boardman, W.S.J.; Lepczyk, C.A. Reply to wolf et al.: Why trap-neuter-return (TNR) is not an ethical
solution for stray cat management. Animals 2020,10, 1525. [CrossRef]
38.
Wolf, P.J.; Rand, J.; Swarbrick, H.; Spehar, D.D.; Norris, J. Reply to Crawford et al.: Why trap-neuter-return (TNR) is an ethical
solution for stray cat management. Animals 2019,9, 689. [CrossRef]
39.
Wierzbowska, I.A.; Olko, J.; Hedrzak, M.; Crooks, K.R. Free-ranging domestic cats reduce the effective protected area of a Polish
national park. Mamm Biol 2012,77, 204–210. [CrossRef]
40.
Galvis, J.O.A.; Baquero, O.S.; Dias, R.A.; Ferreira, F.; Chiozzotto, E.N.; Grisi-Filho, J.H.H. Monitoring techniques in the capture
and adoption of dogs and cats. Geospat. Health 2015,10, 158–162. [CrossRef]
41.
Gotsis, T. Feral Cats: Do Trap-Neuter-Return Programs Work? NSW Parliamentary Research Service: Sydney, NSW, Australia, 2014.
42.
Ve ˇcerek, V.; Kubesová, K.; Vosláˇrová, E.; Bedáˇnová, I. Rates of death and euthanasia for cats in no-kill shelters in the Czech
Republic. Acta Vet. Brno 2017,86, 109–115. [CrossRef]
43.
Sumner, C.L.; Walker, J.K.; Dale, A.R. The implications of policies on the welfare of free-roaming cats in New Zealand. Animals
2022,12, 237. [CrossRef]
44.
Boone, J.D.; Miller, P.S.; Briggs, J.R.; Benka, V.A.W.; Lawler, D.F.; Slater, M.; Levy, J.K.; Zawistowski, S. A Long-Term Lens:
Cumulative Impacts of free-roaming cat management strategy and intensity on preventable cat mortalities. Front. Vet. Sci.
2019
,6,
238. [CrossRef]
Animals 2022,12, 2301 19 of 21
45.
Calver, M.C.; Fleming, P.A. Evidence for citation networks in studies of free-roaming cats: A case study using literature on
Trap–Neuter–Return (TNR). Animals 2020,10, 993. [CrossRef] [PubMed]
46.
Gunther, I.; Hawlena, H.; Azriel, L.; Gibor, D.; Berke, O.; Klement, E. Reduction of free-roaming cat population requires high-
intensity neutering in spatial contiguity to mitigate compensatory effects. Proc. Natl. Acad. Sci. USA
2022
,119, e2119000119.
[CrossRef] [PubMed]
47.
Van Heezik, Y.; Smyth, A.; Adams, A.; Gordon, J. Do domestic cats impose an unsustainable harvest on urban bird populations?
Biol. Conserv. 2010,143, 121–130. [CrossRef]
48.
Zito, S.; Walker, J.; Gates, C.M.; Dale, A. A preliminary description of companion cat, managed stray cat, and unmanaged stray
cat welfare in Auckland, New Zealand using a 5-component assessment scale. Front. Vet. Sci. 2019,6, 40. [CrossRef]
49.
Morgan, S.A.; Hansen, C.M.; Ross, J.G.; Hickling, G.J.; Ogilvie, S.C.; Paterson, A.M. Urban cat (Felis catus) movement and
predation activity associated with a wetland reserve in New Zealand. Wildl. Res. 2009,36, 574–580. [CrossRef]
50.
Aguilar, G.D.; Farnworth, M.J.; Winder, L. Mapping the stray domestic cat (Felis catus) population in New Zealand: Species
distribution modelling with a climate change scenario and implications for protected areas. Appl. Geogr.
2015
,63, 146–154.
[CrossRef]
51.
Industries, M.F.P. Animal Welfare Matters: New Zealand Animal Welfare Strategy. Available online: https://www.agriculture.
govt.nz/dmsdocument/3963-Animal-Welfare-Matters-New-Zealand-Animal-Welfare-Strategy (accessed on 17 July 2022).
52. Ministry for Primary Industries. Animal Welfare Act 1999; Parliamentary Counsel Office: Wellington, New Zealand, 1999.
53. SPSS. IBM SPSS Statistics for Windows, Version 22.0; IBM Corporation: Armonk, NY, USA, 2016.
54.
Aguilar, G.D.; Farnworth, M.J. Distribution characteristics of unmanaged cat colonies over a 20 year period in Auckland,
New Zealand. Appl. Geogr. 2013,37, 160–167. [CrossRef]
55.
McLeod, L.J.; Driver, A.B.; Bengsen, A.J.; Hine, D.W. Refining online communication strategies for domestic cat management.
Anthrozoos 2017,30, 635–649. [CrossRef]
56.
McLeod, L.J.; Hine, D.W.; Bengsen, A.J. Born to roam? Surveying cat owners in Tasmania, Australia, to identify the drivers and
barriers to cat containment. Prev. Vet. Med. 2015,122, 339–344. [CrossRef]
57.
McLeod, L.J.; Hine, D.W.; Bengsen, A.J.; Driver, A.B. Assessing the impact of different persuasive messages on the intentions and
behaviour of cat owners: A randomised control trial. Prev. Vet. Med. 2017,146, 136–142. [CrossRef]
58. McLeod, L.J.; Hine, D.W.; Driver, A.B. Change the humans first: Principles for improving the management of free-roaming cats.
Animals 2019,9, 555. [CrossRef] [PubMed]
59.
Crowley, S.L.; Cecchetti, M.; McDonald, R.A. Diverse perspectives of cat owners indicate barriers to and opportunities for
managing cat predation of wildlife. Front. Ecol. Environ. 2020,18, 544–549. [CrossRef]
60.
Gunther, I.; Raz, T.; Zor, Y.E.; Bachowski, Y.; Klement, E. Feeders of free-roaming cats: Personal characteristics, feeding practices,
and data on cat health and welfare in an urban setting of Israel. Front. Vet. Sci. 2016,3, 21. [CrossRef] [PubMed]
61.
Luria, B.J.; Levy, J.K.; Lappin, M.R.; Breitschwerdt, E.B.; Legendre, A.M.; Hernandez, J.A.; Gorman, S.P.; Lee, I.T. Prevalence of
infectious diseases in feral cats in Northern Florida. JFMS 2004,6, 287–296. [CrossRef] [PubMed]
62.
Kreisler, R.E.; Cornell, H.N.; Levy, J.K. Decrease in population and increase in welfare of community cats in a twenty-three year
trap-neuter-return program in Key Largo, FL: The ORCAT program. Front. Vet. Sci. 2019,6, 7. [CrossRef] [PubMed]
63.
Gilhofer, E.M.; Windschnurer, I.; Troxler, J.; Heizmann, V. Welfare of feral cats and potential influencing factors. J. Vet. Behav.
2019
,
30, 114–123. [CrossRef]
64.
Gunther, I.; Raz, T.; Berke, O.; Klement, E. Nuisances and welfare of free-roaming cats in urban settings and their association with
cat reproduction. Prev. Vet. Med. 2015,119, 203–210. [CrossRef]
65.
Castro-Prieto, J.; Andrade-Núñez, M.J. Health and ecological aspects of stray cats in old San Juan, Puerto Rico: Baseline
information to develop an effective control program. Puerto Rico Health Sci. J. 2018,37, 110–114.
66.
Zito, S.; Aguilar, G.; Vigeant, S.; Dale, A. Assessment of a targeted trap-neuter-return pilot study in Auckland, New Zealand.
Animals 2018,8, 73. [CrossRef]
67. Mirmovtich, V. Spatial organisation of urban feral cats (Felis catus) in Jerusalem. Wildl. Res. 1995,22, 299–310. [CrossRef]
68.
Nutter, F.B.; Levine, J.F.; Stoskopf, M.K. Reproductive capacity of free-roaming domestic cats and kitten survival rate. J. Am. Vet.
Med. Assoc. 2004,225, 1399–1402. [CrossRef] [PubMed]
69.
Gunther, I.; Finkler, H.; Terkel, J. Demographic differences between urban feeding groups of neutered and sexually intact
free-roaming cats following a trap-neuter-return procedure. J. Am. Vet. Med. Assoc. 2011,238, 1134–1140. [CrossRef]
70.
Hayward, J.J.; Taylor, J.; Rodrigo, A.G. Phylogenetic analysis of feline immunodeficiency virus in feral and companion domestic
cats of New Zealand. J. Virol. 2007,81, 2999–3004. [CrossRef] [PubMed]
71.
Gates, M.C.; Vigeant, S.; Dale, A. Prevalence and risk factors for cats testing positive for feline immunodeficiency virus and feline
leukaemia virus infection in cats entering an animal shelter in New Zealand. N. Z. Vet. J.
2017
,65, 285–291. [CrossRef] [PubMed]
72.
Toribio, J.A.L.M.; Norris, J.M.; White, J.D.; Dhand, N.K.; Hamilton, S.A.; Malik, R. Demographics and husbandry of pet cats living
in Sydney, Australia: Results of cross-sectional survey of pet ownership. JFMS 2009,11, 449–461. [CrossRef]
73.
Gates, M.C.; Hinds, H.J.; Dale, A. Preliminary description of aging cats and dogs presented to a New Zealand first-opinion
veterinary clinic at end-of-life. N. Z. Vet. J. 2017,65, 313–317. [CrossRef]
74.
Angel Alberto Florez, M.; Jorge Alberto Solano, M. Demographic study of the population of dogs and cats domiciled in the
southeastern sector of Bucaramanga, Colombia. Rev. de Investig. Vet. Del Peru 2019,30, 828–835. [CrossRef]
Animals 2022,12, 2301 20 of 21
75.
Bruce, S.J.; Zito, S.; Gates, M.C.; Aguilar, G.; Walker, J.K.; Goldwater, N.; Dale, A. Predation and risk behaviors of free-roaming
owned cats in Auckland, New Zealand via the use of animal-borne cameras. Front. Vet. Sci. 2019,6, 205. [CrossRef]
76.
Finkler, H.; Hatna, E.; Terkel, J. The impact of anthropogenic factors on the behavior, reproduction, management and welfare of
urban, free-roaming cat populations. Anthrozoos 2011,24, 31–49. [CrossRef]
77.
Kretzer, M. Cat Found with Leg ‘Hanging on by a Hair’ Is Latest Victim of TNR. Available online: https://www.peta.org/blog/
cat-found-with-severed-leg-is-latest-victim-of-tnr/ (accessed on 17 July 2022).
78.
PETA. Trap, Neuter, Return and Monitor Programs for Feral Cats: Doing It Right. Available online: https://www.peta.org/issues/
animal-companion-issues/animal-companion-factsheets/trap-neuter-return-monitor-programs-feral-cats-right/ (accessed on
17 July 2022).
79.
Andersen, M.C.; Martin, B.J.; Roemer, G.W. Use of matrix population models to estimate the efficacy of euthanasia versus
trap-neuter-return for management of free-roaming cats. J. Am. Vet. Med. Assoc. 2004,225, 1871–1876. [CrossRef]
80.
Brunt, S.; Solomon, H.; Brown, K.; Davis, A. Feline and Canine Rabies in New York State, USA. Viruses
2021
,13, 450. [CrossRef]
[PubMed]
81.
Cho, Y.; Kim, K.; Kim, M.S.; Lee, I. Application of a high-quality, high- volume trap-neuter-return model of community cats in
Seoul, Korea. PeerJ 2020,8, e8711. [CrossRef] [PubMed]
82.
Patterson, M.B.A.; Jamieson, P. Sterilizing pregnant companion animals: Ethics and law. J. Appl. Anim. Ethics Res.
2021
,3, 153–175.
[CrossRef]
83.
Seo, A.; Ueda, Y.; Tanida, H. Health status of ‘community cats’ living in the tourist area of the old town in Onomichi City, Japan.
J. Appl. Anim. Welf. Sci. 2021,25, 338–354. [CrossRef]
84.
Crawford, H.M.; Fontaine, J.B.; Calver, M.C. Using free adoptions to reduce crowding and euthanasia at cat shelters: An Australian
case study. Animals 2017,7, 92. [CrossRef]
85.
Casey, R.A.; Vandenbussche, S.; Bradshaw, J.W.S.; Roberts, M.A. Reasons for relinquishment and return of domestic cats (Felis
silvestris catus) to rescue shelters in the UK. Anthrozoos 2009,22, 347–358. [CrossRef]
86.
Gates, M.C.; Mancera, K.; Dale, A.; Zito, S. Preliminary analysis of post-adoption outcomes for kittens and adult cats rehomed
through a New Zealand animal shelter. N. Z. Vet. J. 2020,68, 38–45. [CrossRef]
87.
Lohr, C.A.; Cox, L.J.; Lepczyk, C.A. Costs and benefits of Trap-Neuter-Release and euthanasia for removal of urban cats in Oahu,
Hawaii. Conserv. Biol. 2013,27, 64–73. [CrossRef]
88. Lamb, W. Commentary on economic valuations of biodiversity. Ecol. Econ. 2013,89, 170–173. [CrossRef]
89.
Legge, S.; Taggart, P.L.; Dickman, C.; Read, J.L.; Woinarski, J. Cat-dependent diseases cost Australia AU$6 billion per year through
impacts on human health and livestock production. Wildl. Res. 2020,47, 731–746. [CrossRef]
90.
Von Dietze, E.; Gardner, D. Euthanizing wildlife: Experiences and coping strategies among people who conduct euthanasia. Pac.
Conserv. Biol. 2014,20, 28–36. [CrossRef]
91.
Rohlf, V.I. Interventions for occupational stress and compassion fatigue in animal care professionals-a systematic review.
Traumatology 2018,24, 186–192. [CrossRef]
92.
Spehar, D.D.; Wolf, P.J. A case study in citizen science: The effectiveness of a trap-neuter-return program in a Chicago neighbor-
hood. Animals 2018,8, 14. [CrossRef]
93.
Spehar, D.D.; Wolf, P.J. The impact of targeted trap–neuter–return efforts in the San Francisco Bay area. Animals
2020
,10, 2089.
[CrossRef]
94.
Barratt, D.G. Predation by house cats Felis catus (L.) in Canberra, Australia. II.Factors affecting the amount of prey caught and
estimates of the impact on wildlife. Wildl. Res. 1998,25, 475–487. [CrossRef]
95. Seo, A.; Ueda, Y.; Tanida, H. Population Dynamics of Community Cats Living in a Tourist Area of Onomichi City, Japan, before
and after the Trap-Test-Vaccinate-Alter-Return-Monitor Event. J. Appl. Anim. Welf. Sci. 2021. [CrossRef] [PubMed]
96. Hampton, J.O.; Fisher, P.M.; Warburton, B. Reconsidering humaneness. Conserv. Biol. 2020,34, 1107–1113. [CrossRef]
97.
Hampton, J.O.; Warburton, B.; Sandøe, P. Compassionate versus consequentialist conservation. Conserv. Biol.
2019
,33, 751–759.
[CrossRef]
98. Sandøe, P.; Corr, S.; Palmer, C. Companion Animal Ethics; Wiley-Blackwell: Hoboken, NJ, USA, 2015.
99.
Castillo, D.; Clarke, A.L. Trap/neuter/release methods ineffective in controlling domestic cat “colonies” on public lands. Nat.
Areas J 2003,23, 247–253.
100.
Natoli, E.; Maragliano, L.; Cariola, G.; Faini, A.; Bonanni, R.; Cafazzo, S.; Fantini, C. Management of feral domestic cats in the
urban environment of Rome (Italy). Prev. Vet. Med. 2006,77, 180–185. [CrossRef]
101.
Carrete, M.; Sánchez-Zapata, J.A.; Benítez, J.R.; Lobón, M.; Camiña, T.; Lekuona, J.M.; Montelío, E.; Donázar, J.A. The precau-
tionary principle and wind-farm planning: Data scarcity does not imply absence of effects. Biol. Conserv.
2010
,143, 1829–1830.
[CrossRef]
102. Read, J. Among the Pigeons: Why Our Cats Belong Indoors; Wakefield Press: Kent Town, Australia, 2019.
103. ACT Government. ACT Cat Plan 2021–2031. A Plan Developed Under the 2017 ACT Animal Welfare and Management Strategy; ACT
Government: Canberra, Australia, 2021.
104. Welsh, P. Cat neutering: The earlier the better to tackle overpopulation. Vet. Rec. 2018,182, 289–290. [CrossRef] [PubMed]
105.
Crawford, H.M.; Calver, M.C. Attitudes and practices of Australian veterinary professionals and students towards early age
desexing of cats. Animals 2019,9, 2. [CrossRef] [PubMed]
Animals 2022,12, 2301 21 of 21
106.
Porters, N.; de Rooster, H.; Moons, C.P.; Duchateau, L.; Goethals, K.; Bosmans, T.; Polis, I. Prepubertal gonadectomy in cats:
Different injectable anaesthetic combinations and comparison with gonadectomy at traditional age. JFMS
2015
,17, 458–467.
[CrossRef] [PubMed]
107.
Porters, N.; de Rooster, H.; Verschueren, K.; Polis, I.; Moons, C.P.H. Development of behavior in adopted shelter kittens after
gonadectomy performed at an early age or at a traditional age. J. Vet. Behav. 2014,9, 196–206. [CrossRef]
108.
Porters, N.; Polis, I.; Moons, C.P.H.; Van De Maele, I.; Ducatelle, R.; Goethals, K.; Duchateau, L.; De Rooster, H. Relationship
between age at gonadectomy and health problems in kittens adopted from shelters. Vet. Rec. 2015,176, 572. [CrossRef]
109.
Moons, C.P.H.; Valcke, A.; Verschueren, K.; Porters, N.; Polis, I.; de Rooster, H. Effect of early-age gonadectomy on behavior in
adopted shelter kittens—the sequel. J. Vet. Behav. 2018,26, 43–47. [CrossRef]
110.
Marston, L.C.; Bennett, P.C. Admissions of cats to animal welfare shelters in Melbourne, Australia. J. Appl. Anim. Welf. Sci.
2009
,
12, 189–213. [CrossRef]
111.
Johnson, J.; Calver, M. Prevalence of desexed cats in relation to age in a convenience sample of Western Australian cats. Aust. Vet.
J. 2014,92, 226–227. [CrossRef]
112.
Mazeau, L.; Wylie, C.; Boland, L.; Beatty, J.A. A shift towards early-age desexing of cats under veterinary care in Australia. Sci.
Rep. 2021,11, 811. [CrossRef]
113.
Lord, L.K.; Wittum, T.E.; Ferketich, A.K.; Funk, J.A.; Rajala-Schultz, P.J. Search methods that people use to find owners of lost pets.
J. Am. Vet. Med. Assoc. 2007,230, 1835–1840. [CrossRef] [PubMed]
114.
Lord, L.K.; Wittum, T.E.; Ferketich, A.K.; Funk, J.A.; Rajala-Schultz, P.J. Search and identification methods that owners use to find
a lost cat. J. Am. Vet. Med. Assoc. 2007,230, 217–220. [CrossRef] [PubMed]
115.
Cansdale, D. Cat and Dog Euthanasia Rates Slashed as Rescue Culture and ‘Fur Babies’ on the Rise. Available online: https://
www.abc.net.au/news/2019-09-11/how-the-rise-of-rescue-culture-and-fur-babieshas-helped/11486424 (accessed on 1 July 2022).
116.
Ho, J.; Hussain, S.; Sparagano, O. Did the COVID-19 pandemic spark a public interest in pet adoption? Front. Vet. Sci.
2021
,
8, 647308. [CrossRef]
117.
McDonald, J.L.; Farnworth, M.J.; Clements, J. Integrating trap-neuter-return campaigns into a social framework: Developing
long-term positive behavior change toward unowned cats in urban areas. Front. Vet. Sci. 2018,5, 258. [CrossRef] [PubMed]
118.
Farnworth, M.J.; Adams, N.J.; Seksel, K.; Waran, N.K.; Beausoleil, N.J.; Stafford, K.J. Veterinary attitudes towards pre-pubertal
gonadectomy of cats: A comparison of samples from New Zealand, Australia and the United Kingdom. N. Z. Vet. J.
2013
,61,
226–233. [CrossRef]