quantifying species similarity and species diversity … species similarity and... · quantifying...

QUANTIFYING SPECIES SIMILARITY AND SPECIES DIVERSITY OF SMALL MAMMALS IN IPEN AND CLOSED HABITATS

Siti Hasmah Binti Taha

middot

Master of Science 2011

QUANTIFYING SPECIES SIMILARITY AND SPECIES

DIVERSITY OF SMALL MAMMALS IN OPEN AND

CLOSED HABIT A TS PKHIDMAT MAKLUMAT AKADEMIK

111111111 rli~ii 111111111 1000246306

SITI HASMAH BINTI T AHA

A thesis submitted in fulfillment of the requirements for the degree of

Master of Science (Quantitative Ecology)

Faculty of Resource Science and Technology UNIVERSITI MALAYSIA SARA W AK

2010

Declaration

I hereby declare that except as acknowledge in the text the work presented in the thesis is entirely my own work and has not been submitted eitherin part or in whole for another degree or qualification at this or any other university or institutions of higher learning

Siti Hasmah Binti Taha November 2010

DEDICATION

I dedicate my work to my wonderful family friends and mentors and to those who know me and make my life worth living for

11

I

Acknowledgments

There had been wonderful opportunity for me in past two years to learn many things that I never

exposed during my undergraduate studies The statistic ecology is interested scope to discovered

since the lack studies conducted about statistic ecology in Malaysia especially in Borneo itself

Million thanks to my principle supervisor Mr Charlie Justin Mergie Laman (Mf Charlie) who

was willing to give me opportunity and believe in me to conduct a great study Without his

support advice and full of guidance probably I could never manage to complete my master

thesis during these two years studies I would like to thank his kindness by assign me as research

assistant for ten months before I register my master study under his supervision Thousand

thanks to my beloved co-supervisor Professor Dr Mohd Tajuddin Abdullah (Mr T) who never

failed to give me full support guidance and scientific experiences to share with me as I can

finished my master thesis I am so grateful since Mr T becoming my second times co-supervisor

after my degree studies in past 3 years Again thousand thanks to him for sharing his valuable

scientific knowledge and for being a good teacher to me

Thousand thanks to UNIMAS which give me strong financial support This research funding

from this study came from UNIMAS fundamental grant FRGS06(05)6542007(l9) In addition

I would like to thank UNIMAS for the ZAMALAH scholarship given from November 2008

until October 20 I O

I would also like to express my personal gratitude to the foHowing Park Warden of Sarna Jaya

Nature Reserve and Kubah National Park for granting pennission and assisting me in getting my

III

fieldwork at Sarna Jaya Nature Reserve (SJNR) and Kubah National Park (KNP) The pennit

number for this study is No NNC9074 (lV)-29 and No 272009

Individual gratitude expressed to Mr Mike Meredith who conducted a good workshop in

statistical methodology to use for my master thesis BeSS workshop on 24 February 2009 until 5

March 2009 which held at Sarna Jaya Nature Reserve gives me more valuable infonnation about

statistical method and an opportunity to apply some of statistical method in my scope of study

Special thanks to the wonderful laboratory assistants Mr Besar Ketol Mr Raymond Patrick ak

Atet Mr Nasron Ahmad and Mr Mohamad Norazlan Bujang Belly from UNIMAS two

important porters from Serian Mr Kevin ak Egoh and Mr Robin ak Jejin for helping me in

making my fieldwork a successful trip Thanks to all of them because willing to help and stay in

national parks for several months Also thanks to the Faculty of Resource Science and

Technology (FRST) UNIMAS administrative staffs for administrative support

Thanks to my loving parents Mr Taha Usup and Mrs Rabiah Johari my brothers sisters and

the special one Mr Mohd Aliy Muhaimin who gives me full moral support and encouragement

during my two years of master study Also not forgotten to the best friends ever from Department

of Zoology (especially to LKULK member PGs from Level 3 Zoonoses room and Museum

room) who always give me support and constructive criticisms about my studies and special

thanks to my colleague Miss Siti Mariam Jamaluddin who shared with me her knowledge

experience and friendship during our field sampling together I will never forget our strong

bonding and good times together

IV

shy

Abstract

( Research on species similarity and species diversity of small mammals in 10 repetitive sampling

occasions (500 trap nights) was conducted in two different habitats an open and closed habitat

The first sampling were carried out in Sarna Jaya Nature Reserve (SJNR) from April to

September 2008 and second sampling conducted in Kubah National Park (KNP) from February

to June 2009 Closed model was chosen where sampling was conducted for five consecutive trap

days in 10 repetitive sessions each session has about five days to one week rest A hundred of

cage traps were set up and placed in grid system at each study site respectively and markshy

recapture method were used to capture the small mammals l

Sarna Jaya Nature Reserve situated in Kuching Sarawak can be classified as geographically

closed habitat or an island habitat A total of 330 individuals of small mammals been trapped in

5000 trapnights consist of six species namely Calloscuirus notatus C prevostii borneensis

Sundamys muelleri Tupaia tana T minor and Rhinoscuirus lauticaudatlls from three families

(Tupaiidae Scuiridae and Muridae) and from two orders (Scandentia and Rodentia) Species

similarities of small mammals species composition on 10 repetitive sampling occasions from

SJNR were calculated and analysed Jaccard similarity coefficient and Sorensen similarity

coefficient indicated a value of 1 in seven tested sampling occasions (3 vs 4 3 vs 5 4 vs 5 3 vs

9 4 vs 9 5 vs 9 and 2 vs 10) Occasion 7 vs occasion 10 noted high values for Morisita-Hom

similarity coefficient (0991) and 0909 is the highest value for Bray-Curtis similarity coefficient

(occasion 9 vs occasion 10) The differences of value for four similarity coefficient indices may

he influenced by factors like sample size and abundance of most abundant species

v

Kubah National Park is classified as a geographically open habitat for this study A total of 98

small mammals were captured in a total of 5000 trapnights consisting of 10 species namely

Maxomys rajah M surifer M whiteheadi Leopoldamys sabanus Lariscus insignis

Sundascuirus lowi Tupaia tana T dorsalis T glis and Echinosorex gymnlllUs from four

families (Erinaceidae Scuiridae Tupaiidae and Muridae) of orders (Rodentia Scandentia and

Insectivora) Species similarity of small mammals in 10 repetitive sampling occasions showed

differences in term of similarity value for four similarity indices All similarity indices noted 0

value for three tested sampling occasions (1 vs 8 5 vs 8 and 8 vs 9) which indicates that there

are no shared species between two tested sampling occasions Occasion 9 vs occasion 10 noted

high value for Jaccard similarity coefficient (075) and Sorensen similarity coefficient (0857)

0982 is the highest value for Morista-Hom similarity coefficient noted from occasion 4 vs

occasion 5 while two tested sampling occasions (4 vs 10 and 6 vs 10) indicate high value for

Bray-Curtis similarity coefficient (075)

The species diversity of small mammals in Kubah National Park (080) is higher than Sarna Jaya

Nature Reserve (046) In conclusion Sarna Jaya Nature Reserve which classified as small

island area comprise of a smaller number in terms of animal species composition due to limited

resources in various aspects (food shelter area) compared to a much wider area which

surrounded by large forested area (Kubah National Park)

KEY WORDS- small mammals similarity indices diversity indices repetitive sampling open

and closed habitat

vi

Abstrak-Kajian tentang persamaan spesies dan kepelbagaian spesies mamalia kedl dalam 10

ulangan ambilan sampel (500 perangkaplmalam) telah di jalankan di dua habitat yang

berlainan habitat yang terbuka dan habitat yang tertlltup Pengambilan sampel yang pertama

telah di jalankan di Hutan Simpan Sama Jaya (SJNR) pada April hingga September 2008 dan

pengambilan sam pel yang kedua di lakukan di Taman Negara Kubah (KNP) pada Februari

hingga Jun 2009 model tertutup telah di pilih di mana pengambilan sampel dyalankan untuk

lima hari berturut-turnt dalam 10 ulangan ambilan sampel setiap ambilan sampel mempllnyai

masa rehat lima hari ke satll minggu Sebanyak seratus perangkap telah di pasang dan di

letakkan mengikut sistem grid di setiap kawasan kajian dan kaedah tanda-tangkap semula

digunakan untllk menangkap mamalia keci

Hman Simpan Sama Jaya terletak di Kuching Sarawak diklasifikasikan sebagai geograji habitat

yang tertutup juga di kenali sebagai habitat pulau Sebanyak 330 ekor mamalia kedl telah di

perangkap dalam 5000 perangkaplmalam mengandungi enam spesies yang dinamakan sebagai

Calloscuirus notatus C prevostii borneensis Sundamys muelleri Tupaia tana I minor dan

Rhinoscuirlls lauticaudatus daripada tiga lamili (Tupaiidae Scuiridae dan Muridae) dan

daripada dua order (Scandentia dan Rodentia) Persamaan dalam komposisi spesies mamalia

kedl dalam 10 ulangan ambilan sampel dari SJNR telah di kira dan di analisa Persamaan

koefisien Jaccard dan persaman koefisien Sorensen menlll1jukkan nilai 1 dalam tujuh lIjian

ulangan ambilan sampel (3 antara 4 3 antara 5 4 antara 5 3 antara 9 4 antara 9 5 antara 9

dan 2 antaJa 10) Antara peristiwa ke 7 dan ke 10 menunjllkkan nilai bagi persamaan koefiesin

Morisita-Horn yang tinggi (0991) dan 0909 merupakan nilai yang tertinggi bagi persaman

k~efisien Bray-Curtis (peristiwa 9 dan 10) Perbezaan pada nilai untllk empat persamaan

Vll

koefzsien indeks mungkin di perngaruhi oleh beberapa Jaktor seperti saiz sam pel danbanyaknua

spe ies yang amat banyak

Taman Negara Kubah (KNP) diklasifikasikan sebagai geograji habitat yang terbuka untuk kajian

ini Sebanyak 98 ekor mamalia kecil telah berjaya di tangkap dalam sejumlah 500

perangkaplmalam mengandungi 10 spesies yang dinamakan sebagai Maxomys rajah M

surikr M whiteheadi Leopoldamys sabanus Larisclis ins ignis Sundascuirus lowi Tupaia tana

T dorsalis I glis dan Echinosorex gymnurLts daripada empat Jamili (Erinaceidae Scuiridae

Tilpaiidae dan Muridae) daripada tiga order (Rodentia Scandentia dan lnsectivora)

Persamaan spesies mamalia kecil dalam 10 ulangan ambilan sampel menunjukkan perbezaan

dari segi nilai persamaan untuk empat persamaan indeks Semua persamaan inde~

menunjukkan nilai 0 untuk tiga ujian ulangan ambilan sampel (1 antara 8 5 antara 8 dan 8

antara 9) yang mana menunjukkan tidak terdapat perkongsian spesies antara dua ujian ulangan

ambilan sampel Ujian antara peristiwa 9 dan 10 menllnjukkan nilai yang tinggi untuk

persamaan koejisien Jaccard (075) dan persamaan koejisien Sorensen (0857 0982

merupakan nilai yang tertinggi bagi persamaan koejisien Morisita-Horn daripada ambilan

sampel 4 dan 5 manakala dua ujaina ambilan sampel (4 antara 10 dan 6 antara 10)

menunjukkan niali yang tinggi bagi persamaan koejisien Bray-Curtis (075)

Kepelbagaian spesies diveristi bagi mamalia kecil di Taman Negara Kubah (080) adalah tinggi

daripada Hutan Simpan Sama Jaya (SJNR) SecQa kesimpulannya Hutan Simpan Sama Jaya

yang juga diklasifikasi sebagai kawasan pulau yang kecil mengandungi jumlah komposisi

mamalia kecil ynag sedikit disebabkan oleh sumber ynag terhad dari pelbagai aspek (makanan

Vlll

~~--~~~------------------------------------------~~-===-----------------~~----

perlidungan dan kawasan) berbanding dengan Taman Negara Kubah yang mempunyai kawasan

yang [lias yang di kelilingi oleh kawasan yang berhutan

KATA KUNCI- mamalia kedl persamaan indeks diversiti indeks ulangan ambilan sampel

habitat terbuka dan tertutup

IX

Pusat Khidmat M klVMERSm MA ya umatAJcademik

uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables

Chapter 1 Introduction

11 General Introduction

12 Species Similarity

13 Species Diversity

14 Cluster Analysis

15 Closure Concepts and Closed Population Model

151 Closure Concepts

152 Closed Population Model

16 Open Habitat and Closed Habitat

17 Small Mammals

171 Biology and Ecology

172 Related Study on Small Mammals

18 Justification

19 Objectives and Hypothesis

191 Objectives

192 Hypothesis

110 Outline ofThesis

X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites

211 Sarna Jaya Nature Reserve 29

212 Kubah National Park 31

Field Methods

221 Sampling Protocol 33

Repetitive Sampling 35

Processing of Specimens

241 Handling and Identification 36

242 Measurements 36

Data Analysis

251 Relative Abundance 36

252 Species Diversity 37

25 21 Shannon Index 37

252 2 Simpson Index 37

253 Species Similarity 39

2521 Jaccards Similarity Coefficients 39

2522 Sorensens Similarity Coefficients 40

25 23 Morisita-Hom Similarity Coefficients 40

25 24 Bray-Curtis Similarity Coefficients 40

254 Cluster Analysis 41

255 Zar t-test 41

Results

Species Composition 43

311 Sarna Jaya Nature Reserve

3111 Species List 43

3112 Cumulative Graph 45

312 Kubah National Park


Xl

I 3122 Cumulative Graph 49

313 Sarna Jaya Nature Reserve and Kubah National Park






33 Cluster Analysis






35 Zar t-test



36 Previous Study

36l Sarna Jaya Nature Reserve 84


Chapter 4 Discussion

41 General Discussion

411 Species Composition 88

412 Factors Affected Capture Probabilities 91




45 Open Habitat and Closed Habitat 111

Xll

Chapter 5 Conclusions and Recommendations

51 Conclusions 114

52 Recommendation for Future Studies 116

References 117

List of publications 132

List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page

1 Maps of Sarna Jaya Nature Reserve 30

2 Maps of Kubah National Park 32

3 Grid sampling design 34

4 Comparison of total individual caught and number of species of small 46 mammals in Sarna Jaya Nature Reserve

5 Total number of small mammals captured in Sarna Jaya Nature Reserve in 47 overall 10 occasions

6 Comparison of total individual caught and number of speCles of small 50 mammals in Kubah National Park

7 Total number of small mammals captured in Kubah National Park in overall 10 51 occaSlons

8 Comparison number of individuals captured in two different sites (Sarna Jaya 56 Nature Reserve and Kubah National Park)

9 Comparison number of species captured ill two different sites (Sarna Jaya 57 Nature Reserve and Kubah National Park)

10 Species accumulation curve in Sarna Jaya Nature Reserve and Kubah National 58 Park

II Jaccards similarity coefficients dendrogram of small mammals in Sarna Jaya 69 Nature Reserve

12 Sorensens similarity coefficients dendrogram of small mammals in Sarna Jaya 70 Nature Reserve

13 Modified Morisitas similarity dendrogram of small mammals in Sarna Jaya 71 Nature Reserve

14 Bray-Curtis similarity dendrogram dendrogram of small mammals ill Sarna 72 Jaya Nature Reserve

XlV

15 Jaccard s similarity coefficients dendrogram of small mammals III Kubah 73 National Park

16 Sorensens similarity coefficients dendrogram of small mammals III Kubah 74 National Park

17 Modified Morisitas similarity coefficients dendrogram of smaH mammals in 75 Kubah National Park

18 Bray-Curtis similarity coefficients dendrogram of small mammals in Kubah 76 National Park

19 Singletons doubletons unique and duplicates of small mammals in Sarna Jaya 79 Nature Reserve

20 Singletons doubletons umque and duplicates of small mammals in Kubah 82 National Park

xv

List of Tables

Page

1 Total sampling efforts for Sama Jaya Nature Reserve and Kubah National Park 35

2 Species composition of small mammals captured in Sama Jaya Nature Reserve 44

3 Species composition of small mammals captured in Kubah National Park 48

4 Comparison of the number of captures and species counts of small mammals at 53 Sama Jaya Nature Reserve and Kubah National Park by using family

5 Number of individuals and relative abundance for small mammals captured at 54 Sama Jaya Naturee Reserve and Kubah National Park

6 Jaccard similarity coefficient index for small mammals at 10 repetitive 60 sampling occasions in Sama Jaya Nature Reserve

7 Sorensen similarity coefficient index for small mammals at 10 repetitive 6 t sampling occasions in Sama Jaya Nature Reserve

8 Morisita-Horn similarity coefficient index for small mammals at 10 repetitive 62 sampling occasions in Sama Jaya Nature Reserve

9 Bray-Curtis similarity coefficient index for small mammals at 10 repetitive 63 sampling occasions in Sama Jaya Nature Reserve

10 Jaccard similarity coefficient index for small mammals at 10 repetitive 64 sampling occasions in Kubah National Park

11 Sorensen similarity coefficient index for small mammals at 10 repetitive 65 sampling occasions in Kubah National Park

12 Morisita-Horn similarity coefficient index for small mammals at 10 repetitive 66 sampling occasions in Kubah National Park

13 Bray-Curtis similarity coefficient index for small mammals at 10 repetitive 67 sampling occasions in Kubah National Park

14 Species diversity of small mammals at 10 repetitive sampling occasions 10 77 Sama Jaya Nature Reserve (EstimateS)

15 Species diversity of small mammals at 10 repetitive sampling occasions 10 80 Kubah National Park (EstimateS)

XVI

16 Value of Zar t-test for small mammals captured in Sarna Jaya Nature Reserve 83

17 Value of Zar t-test for small mammals captured in Kubah National Park 84

18 Previous studies of small mammals in Sarna Jaya Nature Reserve 86

19 Previous studies of small mammals in Kubah National Park 88

20 Summary number of individuals captured and species composition of small 90 mammals at Sarna Jaya Nature Reserve and Kubah National Park

21 Shannon index and Simpson index for Sarna Jaya Nature Reserve and Kubah 91 National Park

22 Summary of similarity indices for small mammals captured III Sarna Jaya 104 Nature Reserve

23 Summary of similarity indices for small mammals captured in Kubah National 106 Park

XVll

CHAPTER ONE

Introduction


Ecological studies can be divided into two components extensive studies and intensive studies

According to Southwood and Henderson (2000) extensive studies frequently carried out to

present an infonnation on distribution and abundance for supplementary conservation and

management program in which the samples area are much larger than intensive studies Intensive

studies generate infonnation about the spatial pattern of populations and it is often possible to

relay the level of the population to oceanographic or climatic factors (Southwood amp Henderson

2000) In many communities studies ecologists acquire a list of the species composition that

occurs in each of several communities and if the quantitative sampling has been conducted some

measure of the relative abundance of each species (Krebs 1999)

Mostly research which been carried out is more focused on species diversity only and just a little

previous tudy been done to quantify the species similarity in addition to species diversity

Quantifying similarity and diversity of animals in tenn of species composition and abundance

will provide useful information about the population of the animal species at the particular study

sites (Magurran 2004)

Species similarity signify the similarity of each study or sample or sampling unit to each other

while the diversity of each unit to one another been signify as species diversity (Krebs 1999) It

is the intension of this study to address both species similarity and species diversity analysis

simultaneously on repetitive sampling of small mammals done on an open and closed habitat


Krebs (1999) noted that a biological community has an attribute of species diversity and recently

there are many different ways have been suggested for measuring the species diversity Basically

it depends on the objective for the study conducted and the information that we want to see from

the samples collected Diversity measures require an estimate of species importance in the

community (Krebs 1999)

Species diversity for this aspect can be defined as the number of different species in a particular

area (species richness) weighted by some measure of abundance such as number of individuals or

biomass (Magurran 1998) Species evenness is part of the measures of species diversity which is

the relative abundance with which each species is represented in the particular area Species

evenness is a measure of biodiversity which qualifies how equal the community is numerically

Evenness is also a part of species diversity measure which compares the similarity of the

population size of each of the species present

Species richness and species evenness are probably the most frequently used measures of the

total biodiversity of the region (Magurran 1998 2004) Species richness noted that the observed

2

species number increases non-linearly with sampling effort Species richness and evenness are

component of biodiversity that mayor may not be correlated with one another and with pattern of

species abundance Gotelli and Colwell (2001) stated that standardising by the number of

individual collected and standardising by area or sampling effort can lead to the different

conclusions regarding species richness

According to Southwood and Henderson (2000) diversity can be divided into three categories

(a) a diversity- the diversity of species within a community or habitat

(b) f3 diversity- the measure of the rate and extent of change in species along a gradient from

one habitat to others and

(c) y diversity- the richness in species of a range of a habitats in a geographical area which

is a consequence of the a diversity of the habitats together with the extent of the (3 diversity

between them

Shannon index is one of several diversity indices used to measure diversity in categorical data

According to Krebs (1999) Shannon index is used for rare species while Simpson index is used

which emphasize the common species The advantage of Shannon index is it takes into account

the number of species and the evenness of the species index is increased either by having

additional unique species or by having better species evenness The Shannon index is the

measure of infonnation content should be used only on random samples drawn from a large

community in which the total number of species is known (Krebs 1999)

3

Shannon index is the most popular measures of the species diversity which based on infonnation

theory According to Krebs (1999) there are four types of infonnation that might be collected

regarding order in the community

(8) The number of species

(b) The number of individuals in each species

(c) The places occupied by individuals of each species and

(d) The places occupied by the individuals as separate individuals

Simpson index also been used to quantify species diversity of small mammals derived by

Simpson (1943) The basis of this index is the probability of two individuals being conspecifics

Here Simpsons Reciprocal Index (110) is used where the values start 1 ([owest possible

figures) and it represents a community containing only one species to the maximum number

which represent the total number of species caught

According to Lande (1996) the species richness Shannon infonnation and Simpson diversity are

the most commonly used nonparametric measures of species diversity The proportion of the total

diversity found within communities provides a natural measure of similarity among multiple

communities The expected similarity among multiple random samples from the same depends

on the number of samples and on the underlying measures of diversity Measures of species

divers~ play a central role in ecology and conservation biology which giving the number of

species composition and the abundance of animals at the particular area

4

uSiJt hidmal MakJumal Akad~rrlliI UMVERsm MALAYSIA SA WAf(


Krebs (1999) noted that resemblance measure can be expressed either (a) similarity degree of

resemblance or (b) dissimilarity degree of difference Magurran (2004) stated that similarity or

dissimilarity indices provide quantitative bases of assessment in comparing species composition

and biodiversity of two or more assemblages in taxonomic and ecological research Krebs (1999)

stated that we need to use similarity coefficients or similarity indices that give some measure of

the relative abundance of each species in the two samples if we are dealing to quantify the

similarity oftwo such samples or community

Similarity matrix is the opposite concept to the distance matrix The elements of a similarity

matrix measure pairwise similarities of objects The greater similarity of two objects will

produced the high value of the measure Each similarity measures has its own properties and

generally gives different perspectives of the data turning the matter of choice not trivial

(Laurenco et al 2004)

According to Boyce and Ellison (2001) and Krebs (1999) similarity indices can be divided into

two major classes first is those that include joint absences or zero-zero matches (species that are

in neither of the two samples for which a similarity is being computed but are present in at least

one other plot in the study) and second class is those that do not include the zero-zero matches

5

QUANTIFYING SPECIES SIMILARITY AND SPECIES

DIVERSITY OF SMALL MAMMALS IN OPEN AND

CLOSED HABIT A TS PKHIDMAT MAKLUMAT AKADEMIK

111111111 rli~ii 111111111 1000246306

SITI HASMAH BINTI T AHA

A thesis submitted in fulfillment of the requirements for the degree of

Master of Science (Quantitative Ecology)

Faculty of Resource Science and Technology UNIVERSITI MALAYSIA SARA W AK

2010

Declaration



DEDICATION


11

I

Acknowledgments




















III





















IV

shy

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

Declaration



DEDICATION


11

I

Acknowledgments




















III





















IV

shy

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

DEDICATION


11

I

Acknowledgments




















III





















IV

shy

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

I

Acknowledgments




















III





















IV

shy

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5





















IV

shy

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

Abstract





















v




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5




















and closed habitat

vi























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5























Vll






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5






















Vlll

~~--~~~------------------------------------------~~-===-----------------~~----





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5





IX


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5


uuolY lA SAJUWAJ(

Table of Contents

Title Page

Declaration

Dedication

Acknowledgement

Abstract

Abstrak

Table of Contents

List of Figures

List of Tables





14 Cluster Analysis





17 Small Mammals



18 Justification


191 Objectives

192 Hypothesis


X

Page

11

III

IV

VI

Vll

x

XIV

XVI

2

5

10

11

12

12

13

20

26

27

28

28

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

Chapter 2

21

22

23

24

25

Chapter 3

31

Methodology

Study Sites



Field Methods





242 Measurements 36

Data Analysis











255 Zar t-test 41

Results







Xl








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5








33 Cluster Analysis






35 Zar t-test



36 Previous Study











Xll


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5


51 Conclusions 114


References 117


List of Appendices

Appendix A 134

Appendix 8 154

Appendix C 160

Appendix D 162

Appendix E 167

Appendix F 174

Appendix G 177

Xlll

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

List of Figures

Page















XlV







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5







xv

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

List of Tables

Page
















XVI









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5









XVll

CHAPTER ONE

Introduction



































2












between them








3




















4



















5

CHAPTER ONE

Introduction



































2












between them








3




















4



















5




















2












between them








3




















4



















5












between them








3




















4



















5




















4



















5



















5

quantifying species similarity and species diversity … species similarity and... · quantifying...

Documents