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Spring 2018

Movement in Cats Valerie N. Plummer

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MOVEMENT IN CATS

by

VALERIE PLUMMER

(Under the Direction of Michelle Cawthorn)

ABSTRACT

Feral cats (Felis catus) are listed as one of the '100 world's worst invasive alien species'.

There are as many as 70-100 million feral cats in the United States as well as an estimated

117-157 million domestic indoor and outdoor cats. Management efforts include a nonlethal

feeding and sterilization program known as "trap-neuter-release" (TNR) where cats are

surgically sterilized and returned to the environment. Population size and structure,

immigration rates, spay/neuter rates, and data on spatial use all play a role in whether TNR

is a viable management option. This study focuses on population structure and spatial use.

To infer the population structure of a population of campus free-roaming cats at the

individual level I used pairwise maximum likelihood estimates of relatedness and

relationship category (unrelated, half-sib, full-sib, parent-offspring). Home range and

movement patterns of domestic free-roaming indoor/outdoor cats were estimated with

100% and 50% adaptive local convex hull and 100% minimum convex polygon for

comparison with previous studies. No differences in home range were found between sex,

age, and season.

INDEX WORDS: Relatedness, Genetics, Ecology, Felis catus, Home-range, GIS, Thesis,

College of Graduate Studies

MOVEMENT IN CATS

by

VALERIE PLUMMER

B.S., Georgia Southern University, 2012

A Thesis Submitted to the Graduate Faculty of Georgia Southern University in Partial

Fulfillment of the Requirements for the Degree

MASTER OF SCIENCE

STATESBORO, GEORGIA

2018

VALERIE PLUMMER

All Rights Reserved

1

MOVEMENT IN CATS

by

VALERIE PLUMMER

Major Professor: Michelle Cawthorn

Committee: C. Ray Chandler

J. Scott Harrison

Electronic Version Approved:

May 2018

2

TABLE OF CONTENTS

LIST OF TABLES .............................................................................................................. 3

LIST OF FIGURES ............................................................................................................ 4

CHAPTER 1 ....................................................................................................................... 5

CHAPTER 2 ..................................................................................................................... 14

Field Methods ........................................................................................................ 14

Genetic Analysis .................................................................................................... 14

Data Analysis ......................................................................................................... 15

GPS and harness fitting .......................................................................................... 16

Data processing and home range ........................................................................... 17

CHAPTER 3 ..................................................................................................................... 19

Kin structure and relatedness ................................................................................. 19

Movement Data ...................................................................................................... 22

CHAPTER 4 ..................................................................................................................... 27

Kin Structure and Relatedness ............................................................................... 27

Movement Analysis ............................................................................................... 29

REFERENCES ................................................................................................................. 33

APPENDIX A GENOTYPES OF INDIVIDUAL CATS ACROSS

NINE LOCI ........................................................................................................ 47

APPENDIX B ALLELE FREQUENCIES ........................................................ 48

APPENDIX C PID AND PIDsibs PER LOCUS AND ACROSS ALL LOCI .. 49

APPENDIX D GENETIC VARIATION PER LOCUS AVERAGE ................ 50

APPENDIX E MATRIX OF RELATEDNESS (RELATIONSHIP) ................ 51

APPENDIX F MATRIX OF RELATEDNESS (R) ........................................... 52

APPENDIX G PRIMER CONCENTRATIONS ............................................... 53

3

LIST OF TABLES

Table 1: Sex, Age, and Reproductive Status..................................................................... 24

Table 2: Home Range Estimates ....................................................................................... 24

Table 3: Mean Home Range Estimates by Season ........................................................... 25

Table 4: Total Tracking and Distance Travelled .............................................................. 26

4

LIST OF FIGURES

Figure 1: Trapping and Feeding Sites ........................................................................... 20

Figure 2: Relatedness at Feeding Sits ........................................................................... 20

Figure 3: Proportion of Related Individuals .................................................................. 21

Figure 4: Home-range Indicators of an Adult Male ..................................................... 23

Figure 5: Female Domestic Free-Roaming Cat with Harness....................................... 23

Figure 6: Domestic Free-Roaming Cat Movements by Time ....................................... 25

5

CHAPTER 1

INTRODUCTION

Biodiversity levels are declining due to habitat loss, pollution, overharvesting,

disease, and the spread of invasive species (Wilson, 1992). Approximately 42% of

threatened or endangered species are at risk because of invasive species (Wilcove et al.

1998; Pimentel et al. 2004), and economic damages associated with controlling invasive

species and their effects amount to approximately $120 billion a year in the United States

(Pimentel et al. 2004).

The spread of an invasive species is influenced by landscape structure, dispersal

mode, and reproductive mode (Moody and Mack, 1988); multiple introductions are often

correlated with the success of invasive species (Barrett and Husband, 1990). Multiple

introductions of the Thiarid snail (Melanoides tuberculata) on Martinique showed

increasing adaptive potential for traits with each successful invasion (Facon et al. 2008).

Other common features of successful vertebrate invaders include a close association with

humans, high population numbers, broad diet and an ability to function in a wide range of

physical conditions (Ehrlich, 1989). The direct and indirect effects of invasive species are

generally negative (Eubanks 2001; Preston et al. 2012). Native to the South Pacific and

Australia, predation from the introduced brown tree snake (Boiga irregularis) has led to

extirpations of 10 native bird species, 6 native lizard species, and 2 native bat species on

the island of Guam (Rodda and Savidge, 2007). Arctic foxes (Alopex lagopus) introduced

to the Aleutian archipelago indirectly induced shifts in plant productivity and community

structure by preying on local seabirds. A reduction in nutrient transport from ocean to

land by seabirds affected soil fertility such that the overall plant composition of the

6

environment changed from grasslands to dwarf shrub/forb-dominated ecosystems (Croll

et al. 2005). Studies of Rattus sp. effects on the soil fertility of offshore islands of New

Zealand showed similar indirect effects (Fukami et al. 2006).

Among extant mammal species, 124 (2.6%) have established a self-sustaining

wild population in at least one location outside their natural range and are classified as

successful invaders (Clout and Russell, 2008). Feral domestic animals (horses, sheep,

goats, cats, dogs) with the help of humans are a part of the relatively few mammal species

that have successfully established populations at more than 30 locations around the world

(Long, 2003). One of the most successful mammalian invaders is the domestic cat (Felis

catus). Listed as one of the '100 world's worst invasive alien species' (Lowe et al. 2000)

there are as many as 70-100 million feral cats in the United States (Jessup, 2004) as well

as an estimated 117-157 million domestic indoor and outdoor cats (Dauphine and

Cooper, 2009). Free-roaming cats (both domestic and feral) are efficient predators and

are generally found at densities 10-100 times higher than native predators (200-1,500 cats

km2: Liberg et al. 2000; Baker et al. 2008) leading to direct impacts on native predators

and native species.

Feral free-roaming cats may survive as lone individuals, or they may be part of

“cat colonies”. Typically, cat colonies are considered to be a group of three or more

individuals living and feeding in close proximity to one another (Slater, 2004). Colonies

may be fully managed (regularly fed, given vaccines, and the population is

spayed/neutered) or semi-managed (regularly fed by a human caretaker), or completely

unmanaged. According to Toukhsati et al., (2007), the persistence of many cat colonies is

likely due to feeding by humans. However, other than feeding no other veterinary care

7

such as spaying and neutering is provided (“semi-ownership” behavior) and thus, the

number of individuals in the colonies varies over time. The breeding system of Felis

catus is conducive to living in a colony as Felis catus is promiscuous (Wilkins, 2007).

Maternal care can either be done individually or communally with the aid of other

mothers in the colony or non-breeder helper cats who provide assistance with grooming

and playing with kittens, guarding the nest or providing food to the mother or kittens

(Kerby, 1988). Females give birth to two to ten kittens per litter with most mating

occurring during the spring and summer due to increasing daylight (Hildreth et al. 2010),

and females can produce up to five litters a year.

All free-roaming cats, including domestic pets, prey on a wide assortment of

native birds (Fitzgerald, 1988), mammals, reptiles, and amphibians (Dunn and Tessaglia,

1994; Crooks and Soule, 1999; Baker et al. 2008). Domestic cats and feral cat diets are

similar with hunting occurring in both domestic and feral free-roaming cats even though

they are given food rations due to natural hunting instincts (Liberg, 1984). Free-roaming

cats are estimated to predate 1.3-4.0 billion birds and 6.3-22.3 billion mammals in the

United States annually (Loss et al. 2013). In addition to the loss of prey species due to

hunting, cats can cause indirect effects on reproductive success of nearby species. The

presence of a free-roaming cat at avian nesting sites reduced feeding rates and induced a

lethal “trait-mediated indirect” effect (Bonnington et al. 2013). Giving up densities and

overall costs of foraging for North American tree squirrels (Sciurus sp., fox/gray) were

higher with the persistent presence of cats and dogs (van der Merwe et al. 2007). Cat

presence also altered the foraging behavior of the dusky hopping-mouse (Notomys

fuscus) more than the dingo (Canis dingo) by reducing patch use (Gordon et al. 2015).

8

Prolific reproduction with early reproductive maturity (as early as six months of age),

relative ease of habitat acclimation, and an innate desire to hunt all make the free-

roaming cat a successful invasive species.

Oceanic islands have provided many examples of the negative ecological impacts

of free-roaming cats (Nogales et al. 2004). Australia has seen a high rate of native

mammal decline. Approximately 33% of all mammals and about 90% of medium-sized

mammals are either extinct or have seen range contractions in Australia (Burbidge and

McKenzie, 1989) and feral cats are widely seen as a significant threat to native fauna and

native fauna conservation. In New Zealand, the extinction of the Stephen’s Island Wren

(Xenicus lyalli) is widely attributed to feral cats. The species was never observed in the

wild and most of the museum specimens collected came from a single cat (Dickman

1996; Fuller 2001).

Previous studies of cat predation have used homeowner reports of prey type and

frequency (Lepczyk et al. 2003; Woods et al. 2003; Baker et al. 2008). However, Lloyd et

al. (2013) quantified domestic cat predation using collar cameras and found that less than

a quarter of cats brought back prey to their residence, suggesting previous homeowner

reports had vastly underestimated the totality of cat predation. Thus, detailed

observations of cats in the field and a description of what and how many prey they kill

are essential to understanding the extent of predatory behavior among free-roaming cats

(Woods et al. 2003).

As our understanding of the overall effects free-roaming cats exert on the

environment and other organisms increases there has been a push for the improvement of

management protocols for cat populations in communities across the United States and

9

worldwide. Management efforts through both lethal (shooting, trapping, poisoning,

hunting) and non-lethal (biological control, exclusion fencing) methods have all been

used in Australia as the native fauna are imperiled (Biodiversity Group Environment

Australia, 1999). However, these methods are seen as expensive and time-consuming

endeavors (Environment Australia, 2001). Lethal methods such as "trap and euthanize"

have been met with opposition from various animal rights groups and the public (Perry

and Perry, 2008). Feline panleukopenia, a virus that primarily attacks the gastrointestinal

system, has been introduced into feral populations on the islands of Jarvis and Marion

and may be effective according to models (Nogales et al. 2004). Poisoning through oral

grooming could potentially be used in the near future (Read, 2010). After many failed

attempts at using a fox/dog baiting strategy, a highly effective baiting technique using

poison laced sausages showed an increase in mortality rates close after bait application

and has been used to eradicate feral cats from two islands off of the Australian coast

(Algar and Burrows, 2004). In the short term, lethal methods tend to rapidly depopulate

cat colonies but they rarely prove to be effective in the long term when colonies are

located on mainland areas (Nutter, 2005). A constant supply of cats from ill-informed

owners ensures complete eradication is nearly impossible.

Non-lethal management methods include a feeding and sterilization program

known as "trap-neuter-release" (TNR) where cats are surgically sterilized and returned to

the environment (Dauphine and Cooper, 2009). Using mathematical models, Andersen et

al. (2004) reported effective population control with the neutering of more than 75% of

the population or the removal of at least 50% of the population. Wallace and Levy,

10

(2006) suggest more ovariohysterectomies than castrations should be performed due to

the predominance of intact females being caught in TNR programs in the United States.

Many factors determine whether TNR will work in a specific circumstance.

Population size and structure, immigration rates, spay/neuter rates, and data on spatial-

use all play a role in whether TNR is a viable option for a group of cats. Feral cat

management according to Stoskopf (2004) should not be a “one size fits all” concept

especially since cats are such an adaptable species. This suggests the most successful

management programs will collect data about the factors presented above, especially

population size and structure prior to implementing a management protocol.

Characterizations of population structure of a species can be useful in many

contexts and is important for understanding the effects of various processes on

populations (Millar and Libby, 1991). Molecular methods of identifying population

structure can be a tool for the management and ecology of wildlife, especially when

paired with behavioral, demographic, or spatial information (DeYoung and Honeycutt,

2005). Identifying population structure through molecular means is valuable when trying

to define significant units of management and conservation (Moritz et al. 1996)

especially for species that are elusive, migratory, or continuously distributed (DeYoung

and Honeycutt, 2005).

Multiple factors can influence allele frequency and diversity including

bottlenecks, founders events, migration, and population size fluctuations. A decrease in

genetic variability is expected to be related to a decrease of population size (Chakraborty

and Nei, 1977). There is large variability in mating tactics within and between

11

populations of the same species (Lott, 1991) and mating systems and dispersal both have

impacts on genetic variability within populations.

Microsatellite loci, a specific sequence of DNA base repeats, have been used in

the analysis of natural populations and provide a greater understanding of population

structure (Slatkin, 1995). They can also be used to identify population attributes such as

relatedness for sensitive or elusive animals (DeYoung and Honeycutt, 2005). A relatively

easy and minimally invasive method to acquire DNA samples is through hair collection

where samples can be directly obtained from a mammal (Higuchi et al. 1988), or

collected as shed hair (Morin et al. 1993). Another source of DNA are fecal samples

(Hoss et al. 1992). While more DNA is found in fecal samples (Broquet et al. 2007),

fresh hair provides higher quality DNA samples versus the other methods (Valderrama et

al. 1999). Hair samples have been used for molecular studies of a wide array of animals

including grizzly bears (Mowat and Strobeck, 2000), black bears (Boerson et al. 2003),

American martens (Pauli et al. 2008), chimpanzees (Morin et al. 1993) and feral cats

(Anile et al. 2012).

Suburban and campus environments serve as habitats to a wide array of

mammals, reptiles, and amphibians as well as resident and migratory songbirds and birds

of prey (Crooks, 2002) and they support local and regional diversity (Angold et al. 2006;

Tratalos et al. 2007). Concerns about depredation of wildlife by free-roaming cats should

extend to these environments (Longcore et al. 2009) because suburban and campus

environments contain fragmented “islands” of natural habitats surrounded by roads and

development (Crooks 2002) that can act as barriers to wildlife movement. This is

especially important because cat abundance generally increases with increasing habitat

12

fragmentation (Crooks 2002), and cat predation is particularly influential in insular

habitats (Nogales et al. 2004). Abundant food sources, like dump and feeding sites, tend

to increase cat survival and population density and subsidized colonies may serve as

source populations for surrounding areas (Schmidt et al. 2007).

Georgia Southern University has an issue with feral cat management. To date,

management decisions have been primarily led by disjointed public efforts. Data on the

current population structure of these cats are unknown and population estimates vary

widely from 20 to 40 individuals. There are several discrete feeding stations scattered

throughout campus that have varying degrees of management. Feeding stations,

dumpsters near academic buildings, and drains tend to have a higher number of cats

present. The spatial ecology of these animals also needs further investigation and how

individuals occupy and use the campus environment specifically is largely unclear.

This study aims to build upon what is known about both feral and domestic free-

roaming cat behavior and biology by investigating the relatedness of feral free-roaming

individuals and the movement and spatial-use of domestic free-roaming cats.

First, I studied a population of feral free-roaming cats (n=24) to obtain data on the

degree of relatedness between individuals. In order to assess the population structure of

free-roaming cats in this area, I calculated pairwise relatedness values and the likelihood

of two individuals being first-ordered relatives (full-siblings, parent-offspring), second-

order relatives (half-siblings) or unrelated to address these questions: 1) What is the

relatedness of adult males in this population? 2) What is the relatedness of adult females

in this population? And 3) are related individuals utilizing the same feeding stations?

Since free-roaming cats exhibit male-biased dispersal and sociality in free-roaming cats is

13

based on females, their relatives, and kittens (Turner, 2000), I predict males will have a

lower degree of relatedness relative to adult females. Social groups in free-roaming cats

form around abundant and rich resources like feeding stations (Macdonald et al. 1987;

Mirmovitch, 1995; Liberg et al. 2000) and individuals are more likely to tolerate and

share resources with kin (Hamilton, 1964) therefore I predict to see kin structure at

feeding stations.

Second, I estimated the home ranges and movement patterns of owned free-

roaming cats in order to gain a better understanding of how cats use and occupy suburban

environments. With GPS tracking data, I aim to address these questions about the spatial-

use of domestic cats: 1) Do home range sizes vary by sex? 2) Does season affect home

range size? And 3) Do domestic cats move more diurnally or nocturnally? I predict home

ranges will vary by sex with males having a larger home range due to male biased

dispersal (Turner, 2000) and I predict a larger home range size in warmer seasons with

more movement occurring nocturnally.

14

CHAPTER 2

METHODS

Field Methods

I studied a population of free-roaming cats located in Statesboro, Georgia

(32.4453° N, 81.7792° W) (Figure 1). I trapped cats 2 to 3 days a week from dusk until

sunrise from March 2014 - January 2016 (IACUC Protocol I13011/I16008). Using a

humane trap (large 32”x10”x12” and extra-large 42”x15”x15” Havahart traps,

Woodstream Corporation, Litiiz, PA) I baited traps with sardines and cat food. A total of

27 cats were captured for this study. Trapping locations were selected prior to trapping

via camera trapping and visual confirmation of the presence of cats. After low trap

success, I deployed a targeted trapping approach at known feeding stations (Figure 1). To

prevent unnecessary mortality, I was present when any trap was deployed and the total

time in the trap for the cat never exceeded 30 minutes. Approximate age (adult, juvenile,

kitten), physical description, and sex (if possible) were recorded at the time of capture.

Wearing gloves, I collected approximately 20 hairs with forceps through the trap to

maximize hair follicle collection for analysis. All samples were placed in small, labeled

envelopes and stored at room temperature at Georgia Southern University.

Genetic Analysis

To maximize DNA extraction hair follicles in each sample were separated from

the shaft using a disposable razor blade. DNA was extracted from 5-10 hair follicles

using a Qiagen DNeasy Blood and Tissue kit following the suggested protocol of the

manufacturer. I used polymerase chain reaction (PCR) to amplify DNA at 10

microsatellite loci that are polymorphic in domestic cats. Primers used for this study

15

(FCA 733, FCA 723, FCA 731, FCA 441, FCA 736, F 124, SRY, FCA 742, FCA 740,

and F 85) were selected from a forensic panel (Menotti-Raymond, 2005). Amplifications

were performed in 20 uL reaction volumes containing 6 uL of DNA, 4 uL cat primer mix,

and 10 uL of 2X Apex Taq Polymerase Master Mix with final reaction concentrations of:

1.5 mM magnesium chloride (MgCl2), 200 uM of four deoxyribonucleoside 5’-

triphosphates (dATP, dCTP, dGTP, and dUTP), 2.0 units of DNA polymerase, and 0.16

mg/mL bovine serum albumin. In order to allow for multiplexing, the markers were

labeled with blue (6-FAM) (FCA 733, FCA 723, FCA 731), green (VIC) (FCA 441, FCA

736, F 124, SRY), and yellow (NED) (FCA 742, FCA 740, F 85) fluorescent dyes at the

five prime end of the reverse or forward primers distinguishing alleles and loci that

overlapped in size. Thermal cycling conditions were taken from Menotti-Raymond et al.

2005 and performed with the GeneAmp 9700. PCR products were visualized on a 2%

agarose gel.

PCR products were combined with an internal size standard and fragment analysis

was performed on an ABI 3500 Genetic Analyzer (Applied Biosystems). Genotypes were

scored using GeneMapper software v4.0 (Applied Biosystems). Alleles were binned by

hand and checked for null alleles and scoring errors with Micro-Checker (Van Ooserhout

et al. 2004).

Data Analysis

To assess the differentiation power of the 10 marker panel (without the SRY

marker) the probability of identity (PID) was calculated (Waits et al. 2001). The

probability of identity with the presence of siblings (PID-sib), defined as the probability

that two randomly drawn sibling individuals from a population have the same multilocus

16

genotype, was also calculated. PID-sib is a conservative upper limit of the possible

ranges for observing identical multilocus genotypes since PID estimates tend to be lower

than observed PID in wildlife populations (Waits et al. 2001).

To determine the likelihood individuals are related I used the program ML-Relate

(Kalinowski et al. 2006). The program calculates maximum likelihood estimates of

relatedness (r) and relationship for codominant genetic data like microsatellites or single

nucleotide polymorphisms (SNPs). The coefficient of relatedness is the probability of two

individuals inheriting an allele identical by descent from a common ancestor. Relatedness

values range from 0 to 1. R values greater than 0 indicate increasing relatedness. For

relationship estimates (unrelated, half-sibling, full-sibling, parent-offspring) each pair is

given a likelihood ratio for a null (Ho) vs. an alternative hypothesis (i.e not related vs.

related). Each hypothesis has two variables, Rm (relatedness maternal) and Rp

(relatedness paternal) that define the probability that two individuals (dyads) share an

allele by direct descent from their mother or father. Three alternative hypotheses were

tested: dyads are half siblings, full siblings, or parent-offspring.

I calculated the relatedness of all adult males and all adult females in the study

separately. Kin structure analysis was done at feeding stations where more than 3

individuals were caught (n=3). All relatedness values are presented as the mean ±

standard error.

GPS and harness fitting

Research on domestic cat movement was carried out in Statesboro, GA (32.4453°

N, 81.7792° W) during 2013 – 2016 (IACUC Protocol I13011/I16008). The subjects of

this study were adult house-based domestic cats. Owners in the study area were contacted

17

and asked to participate in the study beginning in December of 2013. A total of eight

spayed/neutered (at least one year prior to the beginning of the study) cats (3 F, 5 M)

were tracked. The mean approximate age of the domestic cats used in this study was 6.3

years (Table 1). After acquiring permission from the cats’ owners, I fitted each animal

with a waterproof GPS (CatTrack, Perthold Engineering, USA) that was attached to a

backpack-style harness (Figure 5). As a safety measure, the harness was designed with a

release clip that unfastened when pulled. The GPS unit weighed 22g and was well below

the 5% weight limit for each cat (Sikes et al. 2011). Cats were given at least 30 minutes

to acclimate to the GPS and harness. A location was recorded in 5-minute intervals. I

chose this fixed time period to investigate fine-scale movement. When a location fix was

unable to be recorded, the unit would not record until a signal was re-established.

Each cat was tracked for up to eight 24-h periods. No cat was tracked multiple

times or within different seasons. For the welfare of the cat, owners monitored their cats

during deployment and if distress was observed the units were removed immediately.

Data processing and home range

Data recorded on the GPS data loggers included the date, time, longitude, latitude,

altitude, speed, direction, and distance moved by the cat. Before data analysis in ArcGIS

(Version 10.3) location points from the first 30 minutes were removed to account for

harness fitting and re-adjustment time. To define significant movement, the average

distance between two movement points was calculated and any distance below each cat’s

average distance was removed. I estimated home ranges using 100% and 50% adaptive

local convex hull (aLoCoH). This nonparametric kernel method has been shown to be a

better fit for home range estimation due to its ability to identify boundaries in the

18

environment and for convergence to the true distribution as sample size increases (Getz et

al. 2007). For comparison with previous studies, home range was also estimated using

100% minimum convex polygons (MCP) calculated using ArcGIS software. Points were

classified as occurring diurnally or nocturnally by the sunrise and sunset time each day.

Total distance traveled per day and total mean distance traveled per tagging period were

also calculated.

To normalize data, home range estimates were log10 transformed. A t-test was

used to test for differences in home range size for each of the estimators (aLoCoH, MCP)

by sex. The effects of sex and age on home range size were evaluated using an

ANCOVA. Seasonal effects on home range size were calculated using the Kruskal-Wallis

test. Diurnal and nocturnal movement data met normality and equal variance assumptions

and were tested for significance using a paired t-test. All data analyses were done using

JMP (12th edition).

19

CHAPTER 3

RESULTS

Kin structure and relatedness

Out of 27 hair samples collected, DNA was extracted from 24 individuals (20

males, 4 females). The sex ratio of caught individuals in this study is highly skewed to

males. All microsatellites were polymorphic and the number of alleles per locus ranged

from 5-16 (average = 8.667±1.155). Observed and expected heterozygosity was

0.662±0.070 and 0.748±0.048 (ns). The combined probability of identity (PID) using 9

loci is 1.4x10-10 and the PID-sib value is 2.5x10-04. Heterozygote deficiencies were found

in 4 loci (FCA731, FCA 736, F124, and FCA740).

I assessed relatedness at feeding stations where more than 3 individuals were

trapped (Math/Physics, COBA, and Marvin). Mean relatedness at the Math/Physics

(n=5), COBA (n=5) and Marvin (n=5) feeding stations were 0.102±0.037, 0.103±0.042

and 0.338±0.056 respectively. In this population (n=24) overall, adult males had a mean

relatedness of 0.062 ±0.029 while females had a mean relatedness of 0.065±0.065.

Unrelated individuals made up 68% of the individuals found at feeding stations,

followed by 17% of first-order relatives (full siblings, parent-offspring) and 14% of

second-order relatives (half siblings). First-order relatives had a mean relatedness of

0.44±0.03 ranging from 0.338 to 0.6925. Second-order relatives had a mean relatedness

of 0.27±0.04 and relatedness values ranged from 0.1467 to 0.397. Unrelated individuals

had a mean relatedness of 0.063±0.014 with relatedness values ranging from 0 to 0.3257.

No parent-offspring pairs were found at any of the feeding sites.

20

Figure 1 - Trapping and feeding sites located around Georgia Southern University in Statesboro, Georgia.

Circled stars indicate the three main feeding stations.

Figure 2 – Relatedness values at the three feeding stations sampled.

0

0.1

0.2

0.3

0.4

Math/Physics COBA Marvin

Rel

ated

nes

s (r

)

Feeding Station

n=5

n=5

n=5

n=5

n=5

n=5

n=5

n=5

n=5

21

a)

b)

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

First Order Second Order Unrelated

Pro

po

rtio

n o

f R

elat

ed D

yad

s

Degree of Relationship

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

First Order Second Order Unrelated

Pro

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rtio

n o

f R

elat

ed D

yad

s

Degree of Relationship

22

c)

Figure 3 – Proportion of related individuals at a) Math/Physics b) COBA c) Marvin by first-order (full-

siblings, parent-offspring), second-order (half-siblings) and unrelated relationship.

Movement Data

Between 318 and 625 locations were recorded per individual. Over the course of

this study, home range sizes ranged from 2.90 ha to 35.98 ha per cat. No significant

differences were found in home range size by sex (Table 1) for any of the home range

estimators (ns). There were no significant effects of sex or age on home range size

estimators (ns). Since no significant differences between sex or age were found males and

females were combined for seasonal analysis. I found no seasonal difference (Figure 7)

between home ranges (ns) or diurnal and nocturnal movement (ns). Mean distance

traveled per day (Table 3) varied from 1.45 km to 5.12 km with an overall mean of 3.10

km ±0.418 and showed no statistical differences in sex (ns).

0

0.1

0.2

0.3

0.4

0.5

0.6

First Order Second Order Unrelated

Pro

po

rtio

n o

f R

elat

ed D

yad

s

Degree of Relationship

23

Figure 4 – Minimum convex polygon, 50% aLoCoH and 100% aLoCoH of an adult male cat in Statesboro,

Georgia.

Figure 5 – Female domestic free-roaming cat in harness with GPS tracker.

24

Table 1: Sex, reproductive status, and approximate ages of owned free-roaming cats used to determine

home ranges in Bulloch County, GA.

Table 2: Home range estimates by sex (ha; 100% adaptive local convex hull (aLoCoH), 50% aLoCoH,

100% minimum convex polygon) for owned free-roaming cats in Bulloch County, GA. Untransformed

means are presented along with the log10 transformed means and standard error (SE).

Home Range Estimator Sex Mean Mean

logSE

M 7.9021944 0.85121 ±0.16649

F 18.314094 1.0749 ±0.21494

M 0.1735374 -0.8436 ±0.18102

F 0.188509 -0.98578 ±0.23369

M 15.791953 1.13962 ±0.16187

F 34.306826 1.39246 ±0.20897

100% aLoCoH

50% aLoCoH

100% MCP

Home Range (ha)

25

Table 3: Mean home range estimates by season (spring, summer, winter) for domestic free-roaming cats in

Statesboro, GA. a) 100% aLoCoH b) 50% aLoCoH c) 100% MCP.

Figure 6: Domestic free-roaming cat diurnal and nocturnal movement events.

Area (ha)

100% aLoCoH 50% aLoCoH 100 % MCP

Spring 4 9.45 0.18 20.869801

Summer 2 5.35 0.08 16.73

Winter 2 22.97 0.28 32.47

Mean 12.59 0.18 23.35

Season N

0

50

100

150

200

250

300

Diurnal Nocturnal

Nu

mb

er

of

Loca

tio

ns

Time of Day

26

Table 4: Total tracking duration in days, total linear distance traveled in km, and mean distance traveled in

km for each cat.

Cat ID

Total Tracking

Duration

(days)

Total Distance

Travelled

(km)

Mean Distance

Travelled

(km)

1 7 25.84791 3.23098

2 7.8 46.1103 5.12336

3 8 19.88664 2.20962

4 7.4 11.6211 1.45263

5 8.5 38.55547 4.28394

6 6.7 17.57074 2.5101

7 5.2 15.37243 2.56207

8 4 17.44899 3.48979

Mean 6.825 24.0516975 3.10781125

27

CHAPTER 4

DISCUSSION

Kin Structure and Relatedness

Kin structure and dispersal have important consequences for social behavior and

genetic structure (Clutton-Brock and Lukas, 2012). In most promiscuous species of

mammals, males are the predominant dispersers while in monogamous species dispersal

is not skewed towards one sex or the other (Dobson, 1982). Dispersal in feral free-

roaming cats follows the pattern of most mammals where males tend to be the sex that

disperses while females are the philopatric sex (Turner, 2000). Female dispersal,

however, has been previously documented in urban feral cat colonies (Devillard et al.,

2003) where the driving force for mature female dispersal was local resource competition

for quality den sites. When social groups contain more than one breeding female (plural

breeders) and cooperative interactions are common, most females in that group tend to be

related (Sterck et al., 1997) so it is expected free-roaming cats are more related among

females and not males.

Results indicate that adult males in this population had a low mean relatedness.

Low relatedness among males of a promiscuous species is expected due to male-biased

dispersal. Of the 4 females caught only one dyad was related and they were found in

different places on campus (Math/Physics and COBA). Spatial-use data on these two

individuals could potentially reveal shared home ranges. The sampling of more males

than females in this study could be attributed to male-biased dispersal behavior. Further

trapping of individuals, especially females, is needed to accurately assess relatedness in

this population.

28

Two out of the three feeding stations sampled had low relatedness values

(Math/Physics and COBA) with unrelated individuals making up the majority of each

colony. Both Denny et al. 2002 and Shreve, 2014 reported low relatedness values within

their study colonies. The male-biased sex ratio could have contributed to the lower levels

of relatedness observed at these colonies. The Marvin feeding station however had a

higher level of relatedness among individuals caught (0.338±0.056) potentially leading to

more group affiliative behaviors. This feeding station could be a periphery colony since it

is located on the other side of a major road. The road could be a barrier to individual

movement and that could explain the higher observed relatedness values of this feeding

station. If food is dispersed and highly available, like the feeding stations found on and

near campus, cats may not have to compete as fiercely for access and may choose to eat

alone or near relatives (Bradshaw and Hall, 1999).

Previous studies on the effects of TNR on feral cat populations indicates at least

50-75% of the population should be neutered to reduce population numbers and 90% to

offset immigration (Anderson et al. 2004). Some cats are resistant to trapping, as

experienced in this study, and a large amount of abandonment happens on or near the

campus environment making 90% or 100% neutering difficult. In Rome, where there has

been a law mandating the use of TNR programs for a little over 10 years, there has been a

slight decrease in population numbers, but the immigration rate remains high at around

21% (Natoli, 2006), resulting in the necessity of continuing intense efforts of TNR, which

is costly in terms of both economics and time. Without proper education efforts regarding

the behavior and ecology of feral and free-roaming cats and a clear organized campus and

29

public effort unwanted cats will be abandoned negating the results TNR programs may

produce.

Movement Analysis

Much of the research on the spatial ecology of Felis catus and other mammals

utilized very-high-frequency (VHF) radio transmitters. VHF technology requires

receivers to be close enough to triangulate the position of the animal requiring

researchers to be in close proximity, potentially altering animal behavior (Cagnacci et al.

2010). The alternative to VHF tracking used in this study is the use of a global

positioning system (GPS). GPS tracking allows for the collection of animal positions in

remote and poorly accessible areas and during all periods of time and weather conditions

while avoiding altered animal behavior due to researcher proximity (Recio et al. 2011). In

the beginning of GPS development, devices were only suitable for tracking large animals

due to the relatively large receiver and collar. Recently, improvements in battery life and

device size have been developed to track a wider range of animals including hedgehogs,

possums, and feral cats (Recio et al. 2011).

Effective free-roaming cat population control strategies should include a broad

understanding of how they occupy and move through the environment (Bengsen et al.

2012) especially in areas with high conservation value. Spatial information can aid in

predicting the efficiency of different control strategies (Alterio et al. 1999) which include

the method of control, frequency of control efforts, and any costs associated with control.

The majority of studies concerning Felis catus focus on feral free-roaming cat

movement patterns and spatial-use (Liberg 1980; Mirmovitch 1995; Schmidt et al. 2007;

Recio and Seddon, 2013; Kitts-Morgan et al. 2015). Several studies have included

domesticated free-roaming cat home ranges and generally report smaller home range

30

sizes for domestic cats (Schmidt et al. 2007; Horn et al. 2011) than feral free-roaming

cats.

Home range sizes were highly variable among individuals in this study and did

not show statistical differences (Table 2). None of the cats were close enough in

proximity to have overlapping home ranges. The density of cats in the area and habitat

composition may factor into home range determination. In feral free-roaming cats

increasing urbanization and density of cats tend to decrease home range size and if that is

the case for domestic cats further investigation into multiple cat households and

neighborhoods could shed a light on the determinants of home range size and use. This

also poses a management issue since a decrease in cat density could increase overall

home ranges of the cats left in the environment (Thomas et al. 2014) and increase the

probability of mortality and disease.

Home ranges of female cats were not significantly different than male cats in this

study (Table 1) although the youngest female cat in this study had the largest home range

size (35.91 ha) and the largest core home range size (0.46 ha). Younger cats in other

studies, however, tended to have smaller home ranges (Hervias et al., 2014). Reports of

significantly different home ranges vary with some previous studies reporting no

difference in home range size between males and females (Hall et al. 2000; Horn et al.

2007). Other studies indicated males tend to have a larger home range than females

(Liberg 1980; Langham and Porter 1991; Liberg 2000) much like feral free-roaming cats

(Schmidt et al. 2007). Since all the cats were spayed/neutered, reproductive strategies of

space use do not seem to be a driving factor in home range determination in this study.

Domestic free-roaming cats are an interesting case because they are provided with food

31

and shelter and owners residences are a focal point in their home ranges (Thomas et al.

2014). Major roads might be more of a barrier to movement for cats in this area as no

major roads were crossed by any cat in this study.

Future Directions and Conclusions

TNR programs have been implemented on several university campuses in the

United States and abroad with varying success including the University of Central

Florida, Stanford University, Auburn University, Arizona State University, Texas A&M

at College Station and the University of West Georgia. An evaluation of TNR programs

on the University of Central Florida campus found an overall reduction of feral cat

populations over an 11-year period with TNR programs in place (Levy et al. 2003). TNR

programs remain controversial for most campuses. Since feral cats are known to hunt

even with access to supplemental food (Hutchings, 2003) the benefits of feeding stations,

however, are to the cats only and not to reduction population or predation rates (Jones

and Downs, 2011). Feral free-roaming cats are also the definitive host for Toxoplasma

gondii, which is linked to birth defects in humans and they carry other zoonotic diseases

that are of increasing concern to colony caretakers and the community.

The Wildlife Resources Division of the Georgia Department of Natural Resources

has feral cats listed as priority three invasive species falling behind priority one species

like the feral swine (Sus scrofa) and the Coyote (Canis latrans). Nationally, Georgia

ranks 8th in the number of imperiled species and 4th in known or suspected extinctions

(Stein et al. 2000). Sustained monitoring of predation rates and population numbers with

sterilization is needed to determine if the effectiveness of TNR will outweigh its potential

32

costs especially since colony managers tend to underestimate the number of cats present

(Jones and Downs, 2011). Since feral cats are known to hunt even with access to

supplemental food (Hutchings, 2003) the benefits of feeding stations are to the cats only

and not to reduction population or predation rates (Jones and Downs, 2011).

In group-living mammals, individuals are more likely to associate, display

affiliative behaviors, and display less agonistic behaviors with kin over non-kin. Social

organization can influence contact among individuals and the distribution and spread of

disease (Blanchong et al. 2007; Cross et al. 2009). Collecting spatial-use data in

conjunction with social behavior data can help model how diseases move through these

populations (Sanchez, 2012) and aid in management decisions for this controversial

species.

33

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47

APPENDIX A

GENOTYPES OF INDIVIDUAL CATS ACROSS NINE LOCI

KH012 183 183 252 284 342 366 131 135 167 183 292 292 155 159 268 288 318 318

KH016 179 207 284 288 338 342 131 131 167 169 268 268 268 159 244 288 314 318

KH017 179 179 248 272 346 350 127 131 171 171 272 280 123 143 244 264 318 318

KH018 179 207 274 284 342 366 131 131 167 169 268 268 123 159 204 288 318 330

KH019 179 179 248 256 350 366 123 127 183 183 272 304 143 155 272 292 314 318

KH020 179 207 252 272 366 366 119 135 167 201 280 292 147 155 204 288 314 318

KH002 183 207 248 276 342 342 131 131 171 175 292 296 143 143 238 296 318 318

KH21 179 207 244 276 342 370 123 131 183 183 272 276 127 159 276 292 322 330

KH22 179 179 256 272 346 350 127 127 183 183 272 280 123 155 182 272 318 318

KH23 191 207 244 288 346 366 127 131 171 171 268 296 123 143 244 296 314 326

KH24 179 207 268 272 338 346 127 127 171 171 280 296 123 143 244 296 314 318

KH25 179 207 256 268 342 346 127 131 171 171 272 292 123 155 204 268 318 318

KH27 179 207 256 268 342 370 127 135 205 205 292 296 123 147 204 296 318 318

KH28 179 179 256 284 342 342 127 135 167 167 284 296 155 159 266 288 318 318

MA003 179 179 256 262 350 350 131 131 175 175 272 296 147 147 244 244 318 318

MA004 179 179 256 256 350 394 119 131 171 171 272 296 155 163 244 282 338 338

MA005 179 179 262 274 346 394 119 131 175 179 272 296 147 163 188 282 338 338

MA006 179 179 256 256 346 394 119 131 183 201 260 260 147 163 245 245 318 318

MA007 179 179 262 274 346 346 119 131 175 175 272 296 147 163 188 282 318 337

MA009 187 191 256 274 346 346 127 131 179 183 260 284 147 163 244 244 318 318

MA010 179 179 256 256 346 346 119 131 167 179 296 296 147 163 238 244 338 338

MA013 179 179 256 262 346 350 131 131 171 171 272 296 147 163 272 296 318 318

MA014 179 179 272 278 354 354 119 123 169 183 296 296 147 163 262 296 318 338

MA015 179 179 256 262 346 346 131 131 171 179 272 296 147 163 272 296 318 318

FCA 742 F 85 FCA 740

LOCISample

FCA 733 FCA 723 FCA 731 FCA 441 FCA 736 F 124

48

APPENDIX B

ALLELE FREQUENCIES

Locus Allele/n Locus Allele/n Locus Allele/n Locus Allele/n

FCA 733 N 24 FCA 441 N 24 FCA 742 N 24 FCA 740 N 24

179 0.688 119 0.146 123 0.146 314 0.104

183 0.063 123 0.063 127 0.021 318 0.646

187 0.021 127 0.229 143 0.125 322 0.021

191 0.042 131 0.479 147 0.271 326 0.021

207 0.188 135 0.083 155 0.146 330 0.042

FCA 723 N 24 FCA 736 N 24 159 0.104 338 0.167

244 0.042 167 0.146 163 0.188

248 0.063 169 0.063 F 85 N 24

252 0.042 171 0.292 182 0.021

256 0.313 175 0.125 188 0.042

260 0.104 179 0.083 204 0.083

268 0.063 183 0.208 238 0.042

272 0.188 201 0.042 244 0.250

276 0.063 205 0.042 262 0.021

284 0.083 F 124 N 24 264 0.021

288 0.042 260 0.063 266 0.021

FCA 731 N 24 268 0.104 268 0.042

338 0.042 272 0.229 272 0.083

342 0.208 276 0.021 276 0.021

346 0.333 280 0.083 282 0.063

350 0.146 284 0.042 288 0.104

354 0.042 292 0.125 292 0.042

366 0.125 296 0.313 296 0.146

370 0.042 304 0.021

394 0.063

49

APPENDIX C

PID and PIDsibs PER LOCUS AND ACROSS ALL LOCI

Population N FCA 733 FCA 723 FCA 731 FCA 441 FCA 736 F 124 FCA 742 F 85 FCA 740

PID 24 3.0E-01 4.5E-02 6.5E-02 1.4E-01 5.5E-02 6.0E-02 5.5E-02 2.5E-02 2.5E-01

PIDsibs 24 5.8E-01 3.5E-01 3.7E-01 4.4E-01 3.5E-01 3.6E-01 3.5E-01 3.2E-01 5.4E-01

Population N FCA 733 FCA 723 FCA 731 FCA 441 FCA 736 F 124 FCA 742 F 85 FCA 740

PID 24 3.0E-01 1.4E-02 9.0E-04 1.3E-04 6.9E-06 4.1E-07 2.3E-08 5.7E-10 1.4E-10

PIDsibs 24 5.8E-01 2.0E-01 7.4E-02 3.3E-02 1.2E-02 4.2E-03 1.5E-03 4.6E-04 2.5E-04

PID and PIDsibs by Locus

PID and PIDsibs by Increasing Locus Combinations

50

APPENDIX D

GENETIC VARIATION PER LOCUS AVERAGE

51

APPENDIX E

MATRIX OF RELATEDNESS(RELATIONSHIPS)

CAT KH012 KH016 KH017 KH018 KH019 KH020 KH002 KH021 KH022 KH023 KH024 KH025 KH027 KH028 MA003 MA004 MA005 MA006 MA007 MA009 MA010 MA013 MA014 MA015

KH012 -

KH016 HS -

KH017 U U -

KH018 HS FS U -

KH019 U U HS U -

KH020 HS U U HS U -

KH002 HS U U U U U -

KH021 U U U U HS U U -

KH022 U U FS U PO U U U -

KH023 U HS U U U U U U U -

KH024 U U PO U U U U U HS FS -

KH025 HS U U U U U U U U U HS -

KH027 U U U U U U U U U U HS FS -

KH028 HS HS U HS U U U U U U U U U -

MA003 U U U U U U U U U U U U U U -

MA004 U U U U U U U U U U U U U U U -

MA005 U U U U U U U U U U U U U U HS FS -

MA006 U U U U U U U U U U U U U U U HS U -

MA007 U U U U U U U U U U U U U U FS HS FS U -

MA009 U U U U U U U U U U U U U U U U U FS U -

MA010 U U U U U U U U U U U U U U U FS FS HS HS HS -

MA013 U U U U U U U U U U U U U U FS HS U U HS U U -

MA014 U U U U U U U U U U U U U U U U HS U U U U U -

MA015 U U U U U U U U U U U U U U FS U HS U FS U HS FS U -

52

APPENDIX F

MATRIX OF RELATEDNESS (R)

CAT KH012 KH016 KH017 KH018 KH019 KH020 KH002 KH021 KH022 KH023 KH024 KH025 KH027 KH028 MA003 MA004 MA005 MA006 MA007 MA009 MA010 MA013 MA014 MA015

KH012 1

KH016 0.21 1

KH017 0 0 1

KH018 0.25 0.64 0 1

KH019 0 0 0.18 0 1

KH020 0.4 0.05 0 0.18 0.01 1

KH002 0.14 0 0.1 0 0 0 1

KH021 0 0 0 0.08 0.15 0 0 1

KH022 0 0 0.37 0 0.5 0 0 0.07 1

KH023 0 0.18 0.26 0.06 0 0 0.07 0 0 1

KH024 0 0.07 0.52 0 0 0.05 0.06 0 0.16 0.44 1

KH025 0.15 0 0.17 0.03 0 0.01 0.08 0 0.11 0.1 0.23 1

KH027 0.02 0 0 0 0 0.11 0.07 0.14 0 0 0.2 0.4 1

KH028 0.35 0.25 0 0.25 0 0.12 0.11 0 0 0 0 0 0.11 1

MA003 0 0 0.09 0 0 0 0 0 0 0 0 0 0 0 1

MA004 0 0 0.06 0 0 0 0 0 0 0.03 0.03 0.03 0 0 0.13 1

MA005 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.15 0.45 1

MA006 0 0 0 0 0 0.02 0 0 0 0 0 0 0 0 0.14 0.25 0.34 1

MA007 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.28 0.25 0.77 0.18 1

MA009 0 0 0 0 0 0 0 0 0.06 0.04 0 0 0 0.01 0.09 0 0.01 0.4 0.13 1

MA010 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.05 0.35 0.42 0.33 0.34 0.18 1

MA013 0 0 0.29 0 0 0 0 0 0.12 0.03 0.03 0.07 0 0 0.47 0.26 0.18 0.11 0.23 0.05 0.05 1

MA014 0 0 0 0 0 0 0 0.05 0 0 0 0 0 0 0 0.01 0.14 0.05 0.15 0.01 0.2 0.05 1

MA015 0 0 0 0 0 0 0 0 0.03 0 0 0 0 0 0.41 0 0.31 0.11 0.34 0.2 0.3 0.83 0.05 1

53

APPENDIX G

PRIMER CONCENTRATIONS

Loci Dye Primer Concentration Ratio

6-FAM

6-FAM

VIC

VIC

VIC

6-FAM

NED

VIC

NED

NED

0.5

0.5

0.18

0.06

0.025

1

0.81

0.68

0.68

0.68

0.8

0.8

0.3

0.1

0.04

1.6

1.3

1.1

1.1

1.1

FCA733

FCA723

FCA441

FCA736

SRY

FCA731

F85

F124

FCA742

FCA740