appendix 2 – southern cross university genetics report · 2018-10-24 · appendix 2 – southern...

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Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina Koala Plan Prepared for NSW Roads and Maritime Services 54

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Page 1: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

Appendix 2 – Southern Cross University genetics report

Koala Population Viability Analysis – Ballina Koala Plan Prepared for NSW Roads and Maritime Services 54

Page 2: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Page 3: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Page 5: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Page 7: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Page 12: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Alleles Frequencies

Locus Size Repeats WKMP Northern Southern

Phc11 155 15 0.01 - 0.03 157 16 0.21 0.22 0.20 159 17 0.07 0.13 -161 18 0.05 0.04 0.08 165 20 0.05 0.09 -169 22 0.01 0.02 -177 26 0.01 0.02 -179 27 0.17 0.11 0.25 181 28 0.35 0.30 0.43 183 29 0.05 0.07 0.03

Phc13 115 23 0.03 0.06 -117 24 0.14 0.11 0.18 119 25 0.49 0.39 0.63 121 26 0.12 0.19 0.03 123 27 0.13 0.11 0.15 125 28 0.05 0.07 0.03 131 31 0.02 0.04 -133 32 0.01 0.02 -135 33 0.01 0.02 -

Phc25 125 31 0.53 0.50 0.58 129 33 0.07 0.07 0.08 135 36 0.01 0.02 -137 37 0.02 0.04 -147 42 0.16 0.19 0.13 151 44 0.05 0.06 0.05 153 45 0.13 0.09 0.18 155 46 0.02 0.04 -

K2.1 149 13 0.05 0.02 0.10 155 16 0.19 0.28 0.08 159 18 0.27 0.31 0.20 161 19 0.01 0.02 -163 20 0.07 0.09 0.05 173 25 0.36 0.24 0.53 175 26 0.04 0.04 0.05

K10.1 126 10 0.17 0.06 0.33 128 11 0.38 0.31 0.48 130 12 0.03 0.06 -132 13 0.11 0.17 0.03 138 16 0.01 - 0.03 140 17 0.02 0.04 -142 18 0.01 0.02 -146 20 0.10 0.09 0.10 152 23 0.06 0.09 0.03 154 24 0.11 0.17 0.03

3

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Alleles Frequencies

Locus Size Repeats WKMP Northern Southern

Pcv2 121 14 0.01 0.02 -131 19 0.27 0.11 0.28 133 20 0.28 0.26 0.30 135 21 0.32 0.39 0.23 137 22 0.12 0.20 -139 23 0.01 0.02 -

Pcv6.1 211 17 0.05 0.09 -213 18 0.02 0.04 -215 19 0.01 0.02 -217 20 0.20 0.28 0.10 221 22 0.20 0.15 0.28 223 23 0.23 0.30 0.15 225 24 0.28 0.13 0.48

Pcv25.1 72 1 0.01 - 0.03 74 2 0.72 0.67 0.80 82 6 0.05 0.06 0.05 88 9 0.07 0.11 0.03 90 10 0.11 0.11 0.10 94 12 0.03 0.06 -

Pcv25.2 172 174 178

17 18 20

0.85 0.14 0.01

0.81 0.17 0.02

0.90 0.10

-Pcv30 183 47 0.01 0.02 -

195 53 0.45 0.39 0.53 197 54 0.13 0.30 0.03 199 55 0.38 0.33 0.45 201 56 0.02 0.04 -203 57 0.01 0.02 -

Pcv31 213 14 0.40 0.54 0.23 223 19 0.18 0.17 0.20 233 24 0.19 0.04 0.40 239 27 0.22 0.26 0.18

Phci5 146 11 0.09 0.02 0.19 150 12 0.01 0.02 -154 13 0.08 0.06 0.11 158 14 0.14 0.13 0.17 162 15 0.19 0.20 0.17 166 16 0.49 0.57 0.36

Phci9 164 15 0.65 0.61 0.70 167 16 0.14 0.15 0.13 170 17 0.20 0.22 0.18 173 18 0.01 0.02 -

Phci15 208 10 0.10 0.06 0.15 212 11 0.01 0.02 -216 12 0.53 0.41 0.70 220 13 0.35 0.50 0.15 224 14 0.01 0.02 -

4

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WKMP combined Northern Southern

Locus Fis P-value Fis P-value Fis P-value

Phc11 -0.123 0.95 -0.159 0.90 -0.101 0.84

Phc13 0.017 0.14 -0.027 0.17 0.035 0.69

Phc25 0.020 0.09 -0.046 0.10 0.131 0.17

K2.1

K10.1

Pcv2

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Pcv25.1 0.031 0.32 0.031 0.41 0.015 0.58

Pcv25.2 0.015 0.62 0.057 0.55 -0.086 1.00

Pcv30 0.075 0.00* 0.112 0.01* -0.029 0.65

Pcv31 0.002 0.39 -0.186 0.94 0.052 0.45

Phci5 0.046 0.12 0.106 0.03* -0.063 0.58

Phci9 -0.179 0.98 -0.183 0.96 -0.161 0.91

Phci15 0.137 0.27 0.125 0.28 -0.049 0.73

5

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Null Freq. P-value

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Null freq. P-value

Phc11 0 0.95 0 0.90 0 0.83

Phc13

Phc25

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0.216

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0.65

0.15

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K10.1 0.153 0.56 0.203 0.02* 0 0.46

Pcv2 0.036 0.15 0 0.36 0 0.28

Pcv6.1 0.038 0.12 0.063 0.06 0 0.54

Pcv25.1 0.011 0.29 0.019 0.41 0 0.25

Pcv25.2 0.006 0.36 0.021 0.55 0 0.15

Pcv30 0.057 <0.01* 0.072 <0.01* 0 0.50

Pcv31 0.004 0.39 0 0.94 0 0.41

Phci5 0.027 0.09 0.05 0.03* 0 0.47

Phci9 0 0.98 0 0.96 0 0.73

Phci15 0.038 0.25 0.03 0.28 0 0.60

6

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Locus1 Locus2 P-value Locus1 Locus2 P-value Locus1 Locus2 P-value

Phc11 Pcv6.1 0.002 Pcv30 Pcv31 0.003 - - -

Pcv25.2 Phci15 <0.001 Pcv25.2 Phci15 0.003 - - -

7

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n Na ± SE Ne ± SE Pa ± SE Ho ± SE He ± SE F ± SE

Northern 27 6.29 ± 0.5 3.44 ± 0.3 2.14 ± 0.3 0.67 ± 0.04 0.66 ± 0.04 -0.01 ±0.03

Southern 20 4.29 ± 0.4 2.60 ± 0.2 0.14 ± 0.1 0.58 ± 0.04 0.57 ± 0.04 -0.03 ± 0.02

Combined 47 6.43 ± 0.6 3.26 ± 0.3 6.43 ± 0.6 0.63 ± 0.04 0.65 ± 0.04 0.02 ± 0.02

8

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Population Ne 95% CI Ne 95% CI

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Southern 14.9 10.3 – 23.0 8.3 2.5 – 17.5

Combined 47.4 36.8 – 63.3 9.6 5 – 15.7

10

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PARTITION-ML Model Parameters

(df = 111, CV = 87.68 )

k L Χ2

1-null model -3892.43

2 -3833.54 117.78*

3 -3818.39 30.3

4 -3807.20 22.38

5 -3789.24 35.92

15

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Within pops 262 1167.50 4.46 4.46 94

(among ind.) (128) (592.50) (4.63) (0.17) (4)

(within ind.) (134) (575.00) (4.29) (4.29) (90)

Total 267 1253.19 4.74 100

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16

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Population Original pop. Other pop.

Northern 30 1

Southern 97 6

Total 127 7

Percent 95% 5%

17

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Page 63: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

App

endi

x S1

a. M

igra

nts

per

gene

ratio

n (N

m ±

SE)

cal

cula

ted

for

each

of t

he 1

4 lo

ci a

ssum

ing

an e

ffec

tive

popu

latio

n si

ze >

500

for

each

pop

ulat

ion.

A c

utof

f val

ue o

f 0.0

001

was

use

d fo

r S H

UA a

nd a

log

base

=2 t

rans

form

atio

n ap

plie

d fo

r di

ploi

ds. W

-War

della is

res

tric

ted

to t

hose

koa

las

sam

pled

from

fore

st fr

agm

ents

imm

edia

tely

to

the

wes

t of

th

e co

nditi

onal

ly a

ppro

ved

Paci

fic H

ighw

ay u

pgra

de (n

=15)

. W-W

arde

llb incl

udes

all

koal

as s

ampl

ed to

the

wes

t of t

he u

pgra

de (n

=37)

incl

udin

g th

ose

sam

pled

to

the

nort

h an

d no

rthw

est

in a

reas

of t

he f

orm

er B

ig S

crub

rai

nfor

est.

All

othe

r po

pula

tions

are

as

defin

ed in

Nor

man

et

al. (

2015

) an

d A

ppen

dice

s. *

, sig

nific

ance

of

G-t

est

for

alle

le

freq

uenc

y di

ffer

ence

s ca

lcul

ated

for S H

UA w

here

P≤0

.05.

Pop1

E-W

arde

ll

Pop2

W-W

arde

lla

K10.

1

1.53

K2.1

62.1

0

Pcv2

6.93

Pcv2

5.

1 361

Pcv2

5.

2 866

Pcv3

0

28.2

5

Pcv3

1

2.32

*

Pcv6

.1

5.19

Phc1

1

1.47

Phc1

3

4.52

Phc2

5

10.7

7

Phci

15

1.96

*

Phci

5

2.73

Phci

9

-

Nm

± S

E

104

± 69

E-W

arde

ll W

-War

dellb

3.36

74

.61

9.38

83

4 16

32

10.4

9.

83

31.1

4 2.

75

6.34

9.

64

8.96

6.

91

543

227

± 12

7

Nm

bet

wee

n W

arde

ll an

d So

uthe

rn p

opul

atio

ns

War

dell

Sth

Lism

ore

2.97

1.

45*

12.5

4 4.

25*

853

5.09

* 98

.66

0.81

* 1.

30*

3.10

* 0.

52*

21.0

3 6.

03

2954

28

3 ±

214

War

dell

Wes

tern

2.

14*

6.67

7.

70

39.5

1 15

6 6.

18*

136

0.83

* 2.

13*

1.34

* 0.

97*

34.2

9 2.

74*

60.5

7 33

± 1

4

War

dell

Far-

wes

tern

2.

76

0.45

* 1.

07*

4.42

* 53

6 5.

81*

9.96

* 0.

26*

1.73

* 2.

52*

0.84

* 13

.33

1.18

* 15

.99

43 ±

38

War

dell

S.Li

s+W

est

2.35

* 3.

65*

10.9

2 11

.29*

22

1 3.

69*

339

0.66

* 1.

66*

1.82

* 0.

66*

25.7

5 5.

67*

1762

17

1 ±

125

War

dell

All

Sout

hern

6.

06

2.16

* 5.

07*

13.8

6*

591

3.21

* 19

9 0.

47*

1.72

* 3.

29*

0.69

* 27

.15

5.36

* 20

7 76

± 4

4

Nm

bet

wee

n W

arde

ll an

d th

e N

orth

ern

popu

latio

n

War

dell

Nor

ther

n 0.

76*

0.89

* 1.

52*

1.57

* 18

3 2.

92*

2.59

* 0.

24*

3.79

* 3.

35*

0.11

* 1.

32*

1.52

* 9.

01*

15 ±

13

Nm

bet

wee

n W

arde

ll an

d th

e co

mbi

ned

regi

onal

pop

ulat

ions

War

dell

Regi

onal

5.

86*

3.30

* 8.

03*

6.96

* 14

34

8.56

* 23

2 0.

51*

6.95

* 10

.98*

0.

54*

31.7

7*

74.4

5 14

1 14

0 ±

101

4

Page 64: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

Appendix S1b. Effective population size (Ne) for koalas in the WKMP and surrounding areas. Ne calculated using the linkage disequilibrium (LD) and molecular coancestry (MC) methods with 95% confidence intervals (CI). N, sample size. *, denotes sample sizes that are within 10% of the estimated Ne. LD critical values for exclusion of rare alleles in brackets. inf, infinity.

Population n LD (0.02) 95% CI LD (0.01) 95% CI MC 95% CI

Population partitions and Ne for Wardell

E-Wardell 10 148.0 20.3-inf. 148.0 20.3-inf. inf. inf.

W-Wardella 15 21.8 14.9-35.3 21.8 14.9-35.3 inf. inf.

W-Wardellb 37* 37.9 28.8-52.5 20.6 17.1-25.0 12.7 4.4-25.3

Wardell

S Lismore

47* 47.4 36.8-63.6 24.4 20.8-28.9

Population partitions and Ne for regional populations

19 43.6 24.9-123.0 43.6 24.9-123.0

9.6

19.8

5-15.7

0-99.5

Western 23* 19.0 13.7-28.2 19.0 13.7-28.2 2.5 1.4-3.8

S Lis.+Western 42* 73.9 48.8-134.7 41.5 32.4-55.4 4.0 1.8-7.0

Far-western 16 28.6 15.3-92.8 28.6 15.3-92.8 9.0 3.3-17.4

Southern 57 115 74.6-221.8 79.2 58.7-115.6 4.2 2.4-6.4

Northern 31 inf. 193.4- inf. 271.6 98.9- inf. inf. inf.

All Regional 88* 49.7 42.5-58.8 67 56.8-80.3 27.2 7.4-59.7

� �

5

Page 65: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

App

endi

x S1

c. M

igra

nts

per

gene

ratio

n (N

m)

calc

ulat

ed f

or e

ach

of t

he 1

4 lo

ci u

sing

em

piri

cal e

stim

ates

of

effe

ctiv

e po

pula

tion

size

(N

e) c

alcu

late

d by

the

LD

met

hod.

Cu

toff

val

ues

for

S HU

A u

sed

in t

he e

stim

atio

n of

Nm

wer

e se

t at

0.0

001A

(da

sh),

0.00

1B and

0.0

1C , with

tho

se lo

ci f

allin

g be

low

thi

s va

lue

omitt

ed f

rom

the

cal

cula

tion

of

mea

n N

m. W

-War

della is

res

tric

ted

to t

hose

koa

las

sam

pled

fro

m fo

rest

fra

gmen

ts im

med

iate

ly t

o th

e w

est

of t

he c

ondi

tiona

lly a

ppro

ved

Paci

fic H

ighw

ay u

pgra

de (n

=15)

. W

-War

dellb in

clud

es a

ll ko

alas

sam

pled

to

the

wes

t of

the

upg

rade

(n=

37) i

nclu

ding

tho

se s

ampl

ed t

o th

e no

rth

and

nort

hwes

t in

are

as o

f th

e fo

rmer

Big

Scr

ub r

ainf

ores

t.

All

othe

r pop

ulat

ions

are

as

defin

ed in

Nor

man

et a

l. (2

015)

and

App

endi

ces.

Pop1

Po

p2

K10.

1 K2

.1

Pcv2

Pc

v25

.1

Pcv2

5 .2

Pc

v30

Pcv3

1 Pc

v6.

1 Ph

c1

1 Ph

c1

3 Ph

c2

5 Ph

ci1

5 Ph

ci5

Phci

9 AN

m ±

SE

B Nm

± S

E C N

m ±

SE

Nm

bet

wee

n ea

st (E

-) a

nd w

est (

W-)

War

dell

E-W

arde

ll

E-W

arde

ll

W-W

arde

lla

W-W

arde

llb

1.27

4.15

47.3

3

152

4.72

13.3

8

287C

2085

C

675C

909C

28.6

4

20.7

5

1.57

16.7

5

4.97

60.8

7

1.01

2.82

4.14

10.0

4

6.77

7.30

1.63

27.4

0

2.02

12.6

4

-

505C

82 ±

54

273

± 15

6

82 ±

54

273

± 15

6

9.5

± 4.

5

29.8

± 1

3.1

Nm

bet

wee

n W

arde

ll an

d So

uthe

rn p

opul

atio

ns

War

dell

Sth

Lism

ore

1.72

1.

40

10.6

5 2.

54

430C

2.54

69

.42

0.72

0.

83

2.32

0.

41

19.2

4 3.

81

2755

C 23

5 ±

196

235

± 19

6 9.

6 ±

5.7

War

dell

Wes

tern

1.

50

5.37

5.

67

27.2

9 11

0 4.

01

101

0.63

1.

55

0.99

0.

74

25.3

9 2.

01

45.0

4 23

.7 ±

9.9

23

.7 ±

9.9

23

.7 ±

9.9

War

dell

Farw

este

rn

1.31

0.

26

0.38

2.

55

239

2.38

5.

37

0.18

0.

70

1.11

0.

49

4.98

0.

51

8.91

19

.2 ±

16.

9 19

.2 ±

16.

9 19

.2 ±

16.

9

War

dell

S.Li

s+W

est

2.25

4.

67

12.1

8 11

.36

210

3.25

38

2C 0.

74

1.72

1.

98

0.74

29

.22

5.41

20

27C

192

± 14

4 19

2 ±

144

23.6

± 1

7.1

War

dell

Sout

hern

8.

78

4.71

8.

82

20.3

5 70

4C 3.

95

390C

0.88

2.

55

5.43

1.

21

44.9

9 7.

26

410C

115

± 59

11

5 ±

59

9.9

± 3.

7

Nm

bet

wee

n W

arde

ll an

d th

e N

orth

ern

popu

latio

n

War

dell

Nor

ther

n 1.

30

1.97

2.

75

3.26

64

5C 5.

93

4.31

0.

47

5.46

6.

87

0.32

3.

59

3.67

23

.21

51 ±

46

51 ±

46

4.9

± 1.

6

Nm

bet

wee

n W

arde

ll an

d th

e co

mbi

ned

regi

onal

pop

ulat

ions

War

dell

Regi

onal

7.

40

4.71

10

.67

8.83

20

88C

11.0

6 32

6C 0.

69

8.22

14

.26

0.74

47

.06

103

199

202

± 14

7 20

2 ±

147

34.6

± 1

7.1

6

Page 66: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

App

endi

x S1

d. E

stim

ated

dis

pers

al b

etw

een

koal

a po

pula

tions

. Po

int

estim

ates

of

Ne

(Ne-

Pop1

/Ne-

Pop2

) ca

lcul

ated

usi

ng t

he L

D m

etho

d w

ere

used

as

inpu

t ex

cept

tho

se v

alue

s m

arke

d w

ith a

n as

teri

sk (*

) w

hich

cou

ld n

ot b

e re

liabl

y es

timat

ed a

nd t

he lo

wer

CI w

as u

sed.

Mea

n N

m is

giv

en a

s m

igra

nts

per

gene

ratio

n ca

lcul

ated

usi

ng S H

UA

cu

toff

val

ues

of 0

.000

1 an

d 0.

001.

Mig

rant

s/ge

nera

tion

is c

onve

rted

to

mig

rant

s/ye

ar a

ssum

ing

a ge

nera

tion

time

for

koal

as o

f 6.0

2 or

7.8

yea

rs.

Nm

± S

E (S H

UA c

utof

f 0.0

001)

N

m ±

SE

(S HU

A c

utof

f 0.0

1)

Pop1

Po

p2

Ne

(per

gen

.)

(6.0

2 y/

gen.

) (7

.8 y

/gen

.)

(per

gen

.)

(6.0

2y/g

en.)

(7

.8 y

/gen

.)

Mea

n N

m b

etw

een

east

(E-)

and

wes

t (W

-) W

arde

ll

E-W

arde

ll W

-War

della

20*/

22

82 ±

54

13.6

± 9

10

.5 ±

6.9

9.

5 ±

4.5

1.6

± 0.

7 1.

2 ±

0.6

E-W

arde

ll W

-War

dellb

20*/

21

273

± 15

6 45

± 2

6 35

± 2

0 29

.8 ±

13.

1 5

± 2.

2 3.

8 ±

1.7

Mea

n N

m b

etw

een

War

dell

and

Sout

hern

pop

ulat

ions

War

dell

S-Li

smor

e 24

/43

235

± 19

6 39

± 3

2 30

± 2

5 9.

6 ±

5.7

1.6

± 0.

9 1.

2 ±

0.7

War

dell

Wes

tern

24

/19

23.7

± 9

.9

3.9

± 1.

6 3.

0 ±

1.3

23.7

± 9

.9

3.9

± 1.

6 3.

0 ±

1.3

War

dell

Far-

wes

tern

24

/29

19.2

± 1

6.9

3.2

± 2.

8 2.

5 ±

2.2

19.2

± 1

6.9

3.2

± 2.

8 2.

5 ±

2.2

War

dell

S-Li

smor

e+W

este

rn

24/4

2 19

2 ±

144

32 ±

24

24.6

± 1

8.5

23.6

± 1

7.1

3.9

± 2.

8 3.

0 ±

2.2

War

dell

Sout

hern

24

/79

115

± 59

19

.1 ±

9.8

14

.7 ±

7.6

9.

9 ±

3.7

1.6

± 0.

6 1.

3 ±

0.5

Mea

n N

m b

etw

een

War

dell

and

the

Nor

ther

n po

pula

tion

War

dell

Nor

ther

n 24

/99*

51

± 4

6 8.

5 ±

7.6

6.5

± 5.

9 4.

9 ±

1.6

0.8

± 0.

3 0.

6 ±

0.2

Mea

n N

m b

etw

een

War

dell

and

the

com

bine

d re

gion

al p

opul

atio

ns

War

dell

All

regi

onal

24

/67

202

± 14

7 34

± 2

4 26

± 1

9 34

.6 ±

17.

1 5.

7 ±

2.8

4.4

± 2.

2

� �

7

Page 67: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

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Page 68: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

1

PEER REVIEW OF TWO REPORTS:

“AMRI”

“Koala population genetics management. A report to the Roads and Maritime Service (RMS)” 12th August 2015. Linda E. Neaves, Siobhan B. Dennison,, Greta J. Frankham, Mark D. B. Eldridge and Rebecca N. Johnson. Australian Museum Research Institute

and

“SCU”

“Genetic profiling of koalas: Woolgoolga to Ballina Pacific Highway Upgrade (Section 10-Wardell to Coolgardie)”. August 2015. Dr J.A. Norman, Dr C. Blackmore, Assoc. Prof. R. Goldingay & Prof L. Christidis. Southern Cross University.

ASSESSMENT BY: PROFESSOR WILLIAM B SHERWIN Evolution & Ecology Research Centre Deputy Head, School of Biological, Earth and Environmental Science BEES, UNSW Australia, SYDNEY 2052 AUSTRALIA FX: +61 (2) 9385 1558 PH: +61 (2) 9385 2119 EM:[email protected] Web:http://www.bees.unsw.edu.au/staff/academic/Sherwin CRICOS Code: 00098G

UNSW DISLCAIMER

Peer comment is a process of subjecting research methods and findings to the scrutiny of others. The peer comment process is integral to scholarly research. The University of New South Wales (UNSW) attempts to ensure the accuracy and reliability of the information contained in this peer comment. UNSW will endeavour to correct any inaccuracies reported in any information in the peer comment in a timely manner. UNSW accepts no liability for any loss or damage a person suffers because that person has directly or indirectly relied on any information contained in the peer comment.

ASSESSMENT PROVIDED FOR MR SIMON WILSON Environment Officer Environment | Infrastructure Development Roads and Maritime Services Level 2 76 Victoria Street Grafton NSW 2460 T 02 6640 1376 M 0477 315 247 E [email protected] www.rms.nsw.gov.au

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TABLE OF CONTENTS

Page

Title 1

Contents 2

Summary and Recommendations 3

General comments 4

Detailed comments – AMRI report 6

Detailed comments – SCU Report 8

References 10

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SUMMARY and RECOMMENDATIONS

In the Wardell area of northern NSW, a planned upgrade to the Pacific Highway passes through an area inhabited by koalas. NSW Roads and Maritime Services wishes to assess the possible impact of the upgrade on the koala population, and have commissioned two genetic reports: AMRI (by Australian Museum Research Institute) and SCU (by Southern Cross University).

The question “Could the upgrade disrupt koala dispersal sufficiently to reduce the viability of the koala population?” is expressed as a series of deliverables, plus the apparent intention to follow up with a Population Viability Analysis (PVA). I presume that PVA will be used comparatively, examining chance of population persistence over multiple generations, with various levels of dispersal between sub-populations. Such comparative use of PVA is called “sensitivity analysis”.

To set the baseline for such a PVA, genetic methods can be used to assess the natural amount of dispersal between areas. The margins of areas to be investigated can be set for various reasons including habitat suitability or human alteration (eg the highway upgrade).

AMRI and SCU each used a number of different methods to assess genetic subdivision, without converting them to dispersal estimates. It is currently not possible to make a direct comparison between the results of AMRI and SCU at Wardell, because there is insufficient corresponding geographic information. This should be clarified with detailed geographic information for each individual at Wardell and immediate surrounds. However, in both studies, there was a general pattern of increasing genetic similarity at decreasing separation (“Isolation by Disatnce”), but there were no clear boundaries where one locality was genetically isolated from another.

But what is the dispersal across these boundaries? In each report only one method specifically targeted dispersal, by identifying particular koalas who might have dispersed in the most recent generation: those that were genetically assigned to a location other than the one in which they were sampled (AMRI); or those with first-order relatives in a location other than the one in which they were sampled (SCU). Only a small number of individuals were identified as potential dispersers, but it is worth noting that only small numbers are needed to halt genetic differentiation by chance processes in transmission. Thus the areas within Wardell, appear to be important for mutual support of each other, and adjacent areas.

Both studies indicated that dispersal was relatively high both between subpopulations within Wardell, and between Wardell and adjacent areas. This dispersal probably does two things: opposes loss of genetic variation within subpopulations, and forestalls immediate extinction. Therefore, a precautionary conservation management approach would be to avoid any reduction of the measured level of dispersal, pending results of the PVA-sensitivity analysis.

There are two limitations to the use of AMRI’s and SCU’s genetic dispersal estimates as base-line dispersal for the PVA-sensitivity analysis. The first limitation is that sample sizes are unavoidably small, so that the estimates are subject to high error rates. This is compounded by the fact that these particular analyses only tell us about dispersal in the most recent generation. AMRI and SCU were correct to avoid methods that convert their genetic subdivision estimates into dispersal rates, because this has been criticised on various grounds.

However, there exists a Mutual Information method that avoids the problems that beset other genetic dispersal measures, and can deal with the widest possible range of population sizes and dispersal rates. The data from AMRI and SCU should be used in this way, to produce dispersal assessments as a baseline in the PVA-sensitivity analysis, to investigate how the koala population’s viability might be affected if the Pacific Highway upgrade reduces dispersal below this baseline.

AMRI and SCU also produce estimates of Genetic variation within subpopulations, which can be included in some PVA programs.

Finally, in commissioning the analysis of sensitivity of population extinction to altered dispersal, I encourage the Roads and Maritime Services to require information on not only the most likely outcome, but also the worst-case outcome, to facilitate precautionary management.

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GENERAL COMMENTS

In the Wardell area of northern NSW, a planned upgrade to the Pacific Highway passes through an area inhabited by koalas. NSW Roads and Maritime Services wishes to assess the possible impact of the upgrade on the koala population, and have commissioned two genetic reports: AMRI (by Australian Museum Research Institute) and SCU (by Southern Cross University).

The scope of the two reports appears to be slightly different. The question “Could the upgrade disrupt koala dispersal sufficiently to reduce the viability of the koala population?” is expressed as deliverables that are listed by AMRI and SCU. These deliverables are abbreviated as follows.

1. CONNECTIVITY WITHIN WARDELL. AMRI 1: Analyses of population structure and gene flow within the focal area. Also SCU 1: Is the Wardell KMP spatially structured?

2. CONNECTIVITY BETWEEN WARDELL AND SURROUNDS. AMRI 2: Determine whether the focal population appears to be connected to populations in the surrounding area. Also SCU 2: Is the Wardell KMP an important source population for surrounding areas?

3. WITHIN-LOCALITY GENETIC VARIATION. AMRI 3. Allele frequency information for the population as a whole (and for each subpopulation if relevant).

4. VALIDATION. AMRI 4. Summary of procedures undertaken for data validation.

The SCU report indicated the intention to follow these reports with a formal Population Viability Analysis (PVA). I presume that PVA will be used comparatively, examining chance of population persistence over multiple generations, with various levels of dispersal between sub-populations. Such comparative use of PVA is called “sensitivity analysis” (Penn et al. 2000; Reed 2009).

To set the baseline for such a PVA, genetic methods can be used to assess the natural amount of dispersal between areas. The margins of areas to be investigated in this way might be set for various reasons including habitat suitability, human alteration (eg the highway upgrade), or intrinsic differences such as genetic incompatibilities (the latter are unlikely in the case of koalas, Sherwin et al. 2000). In these reports, the AMRI and SCU are asked to assess dispersal between areas within Wardell KMP (deliverable 1), and between Wardell KMP and adjacent areas (deliverable 2).

The spatial arrangement of sampling within Wardell was evident in the SCU study, but not in the AMRI study – perhaps AMRI did not have access to detailed location data.

There are many different methods that use genetic data to estimate average levels of dispersal between areas for a PVA. The AMRI and SCU reports each use a number of different methods to assess genetic subdivision, most of which were not converted to dispersal estimates. In both studies, there was a general pattern of increasing genetic similarity at decreasing separation (“Isolation by Disatnce”), but there were no clear boundaries where one locality was genetically isolated from another.

In each report only one method specifically targeted dispersal, by identifying some individuals who might have dispersed in the most recent generation: koalas that were genetically assigned to a location other than the one in which they were sampled (AMRI); or koalas with first-order relatives in a location other than the one in which they were sampled (SCU).

Both studies indicated that dispersal was relatively high both between subpopulations within Wardell, and between Wardell and adjacent areas. As the authors point out, this dispersal probably does two things: opposes loss of genetic variation within subpopulations (thus potentially aiding future adaptability, Frankham et al. 2010), and forestalls immediate extinction, which is a risk in small isolated populations, including koalas (Lunney et al. 2002).

Therefore, a precautionary conservation management approach would be to avoid any reduction of the measured level of dispersal, unless there had been assessment of the potential effects of such reduction. However, if PVA-sensitivity analysis shows that a certain reduction of dispersal (x%) is not

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likely to significantly affect population viability, then it would be reasonable to implement particular management options, if it could be reliably demonstrated that these options would reduce dispersal by no more than x%.

SCU recommends that the genetic measures of dispersal be incorporated into the future PVA. There are two limitations to the use of AMRI’s and SCU’s genetic dispersal estimates as base-line dispersal for the PVA-sensitivity analysis. The first limitation is that sample sizes are unavoidably small, so that the estimates are subject to high error rates. This is compounded by the fact that these particular analyses only tell us about dispersal in the most recent generation. AMRI and SCU were correct to avoid methods that purport to assess dispersal over tens of generations, but have been criticised on various grounds.

However, there exists a Mutual Information method that can successfully assess dispersal over tens of generations, from genetic data (Sherwin 2006, 2010, Dewar et al 2011, Chao et al 2015). This method avoids the problems that beset other genetic diversity and dispersal measures, and, unlike the other methods, can deal with the widest possible range of population sizes (10 upwards) and dispersal rates (from one in a thousand, to one-third of the population, per generation; Sherwin 2006). All the data from AMRI and SCU should be analysed by the Mutual Information method, to produce robust multigeneration dispersal assessments; the Mutual Information calculations can be carried out in the freeware GENALEX (http://biology.anu.edu.au/GenAlEx) which was used for other tasks by AMRI. The GENALEX website also contains a guide for conversion of Mutual Information for microsatellites to a dispersal estimate. For the mitochondrial DNA, such a conversion could be achieved by following either Dewar (2011, equation 8) for each variable site, or Chao et al. (2015 supplement equations B5-7) for entire haplotypes.

These dispersal assessments from Mutual Information could then be used as baseline in the PVA-sensitivity analysis to investigate how the viability of the koala population might be affected if the upgrade of the Pacific Highway reduces dispersal below this baseline level. The PVA-sensitivity analysis should assess the effect of the highway upgrade, including any measures likely to increase or decrease the road corridor’s permeability to koalas, such as fences or overpasses.

The other data that can be included in the PVA is AMRI’s and SCU’s estimates of genetic variation within subpopulations, which can be included in some PVA programs. Its inclusion will add to realism, especially allowing assessment of when the genetic diversity at Wardell might fall below the lowest levels listed in populations of koalas and other species, reviewed by AMRI.

Finally, in commissioning the analysis of sensitivity of population extinction to altered dispersal, I encourage the Roads and Maritime Services to require information on not only the most likely outcome, but also the worst-case outcome, to facilitate precautionary management.

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DETAILED COMMENT – AMRI REPORT

SAMPLING THE LOCALITIES AND INDIVIDUALS:

From Wardell there were 38 samples for microsatellite and mitochondrial DNA. The lack of identification of position for East and West Wardell makes it difficult to assess their importance relative to the proposed highway upgrade, and to compare this to the SCU report, which uses other nomenclature. Also the tiny number of samples from East makes any conclusions weak.

Four other sites near the NSW/Queensland border (Macquarie, Coffs Harbour, Tyagarah and Coomera) were sampled for both microsatellite and mitochondrial DNA, plus a sample set from the whole koala range for mitochondrial DNA only.

THE GENES USED AND THEIR VALIDATION – DELIVERABLE 4.

1. Microsatellites – biparental inheritance

The analysis was based on a good number of genes -15, and appeared suitable for the tasks of determining variability within location, and differentiation and dispersal between locations. The genes were checked carefully. Two other microsatellite genes were excluded for good reasons, and 10% of individuals were independently re-genotyped. The probability of two individuals having the same microsatellite profile was low (10-15), showing that a good battery of genes had been analysed. Variants at the 15 genes appeared to be inherited independently, so that each provided useful information for the analysis (no “linkage disequilibrium”). In most cases there was no evidence of non-random mating within site (ie, there were few cases of genes out of “HWE”).

2. Mitochondrial DNA – maternal inheritance

An 800 bp portion of the mitochondrial DNA control region was sequenced, and appeared suitable for the tasks of determining variability within location, and differentiation and dispersal between locations.

WITHIN-LOCALITY GENETIC VARIATION – DELIVERABLE 3.

Microsatellite diversity within locations was summarised by a suitable array of measures: allelic diversity, allelic richness, private alleles (AP on page 6, called Pa later in Table 1) expected and observed heterozygosity, Hardy-Weinberg equilibrium (HWE–Fis Table 1) and linkage disequilibrium. These measures were not out of the ordinary for koalas (Table 1).

For mitochondrial DNA, within-locality variation was assessed by suitable statistics - haplotypic diversity and nucleotide diversity - within Wardell and the four other main sites. Wardell values were not out of the usual for koalas, although 37 out of 38 individuals had the same mitochondrial genotype (haplotype) at Wardell.

CONNECTIVITY WITHIN WARDELL _DELIVERABLE 1.

And

CONNECTIVITY BEWTEEN WARDELL AND SURROUNDING AREAS – DELIVERABLE 2.

1. Microsatellites – biparental

Microsatellite geographic structure was assessed by a number of suitable methods: STRUCTURE, DAPC, F-statistics, AMOVA, Isolation-by-distance tests in Mantel, Spatial autocorrelation of pairwise relatedness in GENALEX 6.5. Many analyses were presented without saying which type of gene (microsatellite or mitochondrial) they were based upon; I believe that in all such cases, they were microsatellites.

The authors avoided specifying definitive management units, which I consider to be wise given the relatively low differentiation indicated by most measures. There was only one genetic cluster at Wardell (deliverable 1), and gradually increasing differentiation with distance from Wardell, but no sharp breaks, a pattern called “Isolation by Distance” (deliverable 2).

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As well as the DAPC, there was also a PCA presented on P11 (Fig 3), but not described in the methods section. This appears to be an analysis of microsatellite data, though that is not stated. The PCA showed that the five koalas from “East of focal area” were scattered amongst those from the west of focal area. The text states that the data in Fig 3 come from within Wardell. Thus “East of focal area” appears to mean the East part of Wardell itself, rather than an area to the east of the Wardell area, which would be the interpretation in other parts of the document, where the whole of Wardell appears to be referred to as the “focal area”. Perhaps for the purposes of the PCA, the “focal area” means the proposed upgraded highway. If that interpretation is correct, then there appears to be no justification, at least with this small sample, for considering the koalas on either side of the proposed upgraded highway to be members of distinct separate populations. This should be clarified with detailed geographic information for each individual, so that there could be direct comparison with the results of SCU, which is currently not possible.

There were only two exceptions to the pattern of low differentiation, but I would not prioritise these two findings over the general consensus that there is little geographic differentiation within Wardell or between Wardell and other populations). Fst and Phi-st did show significant departures from zero, but there are many criticisms of Fst, and only partial fixes for these criticisms (Sherwin 2010, Wang 2015). Phi-st likely suffers from many of the same problems as Fst, because Phi-st is also a variance partition and an “order 2” diversity measure (Hill, 1973), the two characteristics that are at the root of Fst’s many problems.

Of course, low differentiation may be due to high dispersal, and some of these measures (including Fst) can be converted to measures of dispersal, but the authors wisely did not do so, given the criticisms mentioned already.

However, there was one assessment of dispersal in and out of Wardell by microsatellites. Microsatellite DNA is biparentally inherited, so it traces dispersal of both sexes. Microsatellites were used to assess dispersal by identifying some individuals who might have dispersed in the most recent generation: koalas that were genetically assigned to a location other than the one in which they were sampled. The assignment test used was in GENALEX 6.5. It showed that some individuals were likely to have moved between Wardell and nearby localities such as Coffs Harbour, but the authors noted that the conclusions were hampered by a lack of samples from localities immediately adjacent to Wardell. Only a small number of individuals were identified as potential dispersers, but it is worth noting that only small numbers are needed to halt genetic differentiation by chance processes in transmission (Kimura and Crow, 1970).

2. Mitochondrial DNA – female dispersal

Mitochondrial DNA generally confirmed the results of the microsatellite analysis, but indicated slightly reduced dispersal of females, relative to males

Mitochondrial DNA is maternally inherited, so traces female dispersal. Mitochondrial geographic structure was assessed by suitable methods: AMOVA/Phi-ST versus distance, and a haplotype network. The AMOVA showed that 92% of mitochondrial variation was within locations. This contrasts with 75% of biparentally-inherited microsatellite variation being within locations, suggesting limited female dispersal. However, Wardell mitochondrial haplotypes do occur elsewhere, so that there must be some female dispersal. Also, mitochondrial DNA suggested that one individual was an immigrant to the Wardell area.

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DETAILED COMMENT – SCU REPORT

SAMPLING THE LOCALITIES AND INDIVIDUALS:

Tables 1 and 5 show 47 samples sourced from the Wardell KMP plus two adjacent localities to the north: Lynwood and Dalwood. This group of samples will be collectively referred to as “Wardell” throughout this assessment. There were also an additional 88 koalas outside Wardell, whose locality information was somewhat scattered in the document, but from Figure 4 it seems that there were three sample sets from localities immediately to the west of Wardell, named from north to south as 30 koalas from “North Lismore”, 20 from “South Lismore” and 22 from “Western”. There were also 16 other koalas from further to the west of Wardell (“Far-western”).

THE GENES USED AND THEIR VALIDATION – DELIVERABLE 4 (not listed as such by SCU).

The SCU analysis used only microsatellite genes - fourteen of them, and adequate number. It is not clear if this set of genes overlaps the set of genes analysed by AMRI. The variation at these genes was sufficient to give a 99% chance that a non-parent would be excluded as a potential parent. Microsatellite analysis of one known parent-offspring pair gave a relatedness estimate of approximately 0.5 (the correct value for such a pair. These values give confidence to the subsequent work assigning first-order relatives (FOR – parent-offspring or full-sibling). It was stated that these microsatellites are “able to detect the presence of genetic differentiation amongst populations with a power of 0.975 or higher after 10 generations and assuming an effective population size of 50-200”. It was not explained how this power analysis was carried out.

WITHIN-LOCALITY GENETIC VARIATION – DELIVERABLE 3 (not listed as such by SCU).

Microsatellite diversity within-locations was not unusual for koalas (Table 5). In the north part of Wardell, there was marginally higher genetic variation than in the south (Table 5, with no confidence limits, so the significance of the difference cannot be evaluated). In the north part of Wardell, there was also lower mean relatedness (Table 3). If real, these two differences could indicate that the north has larger population size, or that it receives more immigration from elsewhere.

CONNECTIVITY WITHIN WARDELL _DELIVERABLE 1.

Genetic Subdivision

There appears to be mild genetic substructure within the Wardell area, but no complete isolation.

On Page 10 it is stated that “Genetic neighbourhood size in the Wardell KMP was estimated to be 21-30 Km2. This confirms a pattern of limited dispersal across the study area and the likely presence of multiple subpopulations.” A genetic neighbourhood is the size of an area within which mating appears to be random. Fig 1 indicates that the Wardell area is about 6km x20km, so that multiple neighbourhoods a few km across could indeed fit into the Wardell area. However, note that neighbourhood calculations are based on the idea that the population is continuous over a much larger scale than the neighbourhood, so they give no indication of sharp boundaries – indeed they assume that no such boundaries exist.

Sharp boundaries were also not supported by the FOR analysis, which suggested that connectivity between localities within Wardell is greatest between the closest localities, and decreases with distance (called “Isolation by Distance” page 10, paragraph 2).

Fst and a related quantity Rst suggest subdivision within Wardell. These measures of genetic differentiation are relatively high between North and South Wardell, compared to their values for differentiation between Wardell and the two closest localities (South Lismore and Western) west. However, these values are presented with neither significance testing, nor confidence limits. I suspect that the latter would be so wide that the comparison is meaningless - Fst has poor statistical properties, as discussed above.

On page 12 it is stated that “We also reject a model in which the Wardell KMP is divided into an eastern and western subpopulation corresponding to the two large tracts of remnant schlerophyll woodland and forest”. It should be clarified why this model was rejected, and where on the map are

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the two large remnant tracts. It is also not clear how to compare this result to AMRI’s “East” and “West”, but if the division is the same in the two reports, then AMRI’s finding in their PCA would confirm the SCU assertion. However, the correspondence cannot be known until the geographic and genetic data for both studies are plotted on a single map.

Genetic Estimates of Dispersal

Only one method specifically targeted dispersal, by identifying some individuals who might have dispersed in the most recent generation: koalas with first-order relatives (FOR) in a location other than the one in which they were sampled. Fig 3 showed that first-order relative pairs were found to be shared between most parts of Wardell , though decreasingly so at greater distances. Most pertinently for the purpose of the study, on pages ii and 10 it is stated that the FOR analysis confirmed that dispersal occurs across the proposed highway upgrade, at two places: Bagotville in the south, as well as in the north.

CONNECTIVITY BEWTEEN WARDELL AND SURROUNDING AREAS – DELIVERABLE 2.

Similarly to the pattern within Wardell, there is some evidence that localities more distant to Wardell are more genetically differentiated from Wardell (Table 4, again without significance testing or confidence limits). This is also reflected in the pattern of inferred dispersal events (Fig 4).

The authors also suggest that dispersal into the northern Wardell area is indicated by its relatively high levels of genetic variation (Table 5), however, as discussed above, the difference is marginal and has no confidence limits to allow assessment of its significance.

It was asserted several times that dispersal was asymmetrical, but few data were available to confirm this. There are programs such as MIGRATE that can attempt to fit models of asymmetric dispersal to genetic data, but I suspect that these programs would fail to converge, due to lack of data. With the existing smaller dataset, it might be possible to infer directionality of dispersal from the FOR data, if (1) there are data on ages of members of each FOR pair, (2) it is assumed that the younger member of the pair is an offspring, and (3) it is assumed that offspring are more likely to disperse. These assumptions mean that such an analysis might have only dubious value.

OTHER

Page 17 talks of “the potential significance of the Southern subpopulation of the Wardell KMP as the remaining relatively pure gene pool for koalas in this region”. It is not clear how genetic purity is defined, nor is it explained why genetic purity is needed. Generally, the opposite - higher genetic variability - is good for conservation management (Frankham et al 2010), unless there are problems of genetic incompatibility between different races of koalas, which no-one has every suggested, to my knowledge.

Page 17 says that the south part of Wardell is more like a “functional koala metapopulation” than the north. There are two definitions of “metapopulation” (Levins 1969, Hanski 1999), either of which could probably apply to both north and south Wardell. I recommend that this term should not be used without further explanation.

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REFERENCES

Dewar RC, Sherwin WB, Thomas E, Holleley CE, Nichols RA. 2011. Predictions of single-nucleotide polymorphism differentiation between two populations in terms of mutual information. Molecular Ecology 20:3156–3166.

Frankham RF Ballou JD & Briscoe DA 2010. Introduction to conservation genetics. Cambridge Univ Press Cambridge.

Hanski, I. Metapopulation Ecology Oxford University Press. 1999

Hill, M.O. Diversity and evenness: a unifying notation and its consequences. Ecology 1973, 54:427-432.

Kimura M and Crow JF 1970. An Introduction to Population genetics Theory. Harper and Row, New York.

Levins, R. 1969. Some demographic and genetic consequences of environmental heterogeneity for biological control. Bulletin of the Entomological Society of America 15: 237–240

Lunney D, O’Neill L, Matthews A and Sherwin WB 2002. Modelling mammalian extinction and forecasting recovery: koalas at Iluka (NSW Australia). Biological Conservation 106:101-113.

Penn AM, Sherwin WB, Gordon G, Lunney D, Melzer A and Lacy RC. 2000 Demographic Forecasting in Koala Conservation. Conservation Biology 14:629-638.

Reed J 2009. Vital rate sensitivity analysis as a tool for assessing management actions for the desert tortoise. Biological Conservation 142:2710–2717.

Sherwin WB, Timms P, Wilcken J, and Houlden B. 2000. Analysis and Conservation Implications of Koala Genetics. Conservation Biology 14:639-649.

Wang J 2015. Does Gst underestimate genetic differentiation from marker data? Molecular Ecology (2015) 24, 3546–3558

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Craig Moritz, FAA Director, Centre for Biodiversity Analysis Professor & ARC Laureate Fellow T: +61 2 6125 5651 or 9492 Evolution, Ecology & Genetics F: +61 2 6125 5573 Research School of Biology E: [email protected] Building 116 Canberra ACT 0200 Australia�<@���%)�"��'�<:;?��

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Page 79: Appendix 2 – Southern Cross University genetics report · 2018-10-24 · Appendix 2 – Southern Cross University genetics report Koala Population Viability Analysis – Ballina

_________________________________________________________________________________________________________________________________ ANU COLLEGE OF MEDICINE, BIOLOGY & ENVIRONMENT

Craig Moritz, FAA Director, Centre for Biodiversity Analysis Professor & ARC Laureate Fellow T: +61 2 6125 5651 or 9492 Evolution, Ecology & Genetics F: +61 2 6125 5573 Research School of Biology E: [email protected] Building 116 Canberra ACT 0200 Australia%�))�'#0��(��(�)�����#�'4(��!��(�"%!�#����'$((�)���'���$#3�����$'��#�!.0�����#��)���'�(*!)(�$���$'"�#��)��!3�"$'��'�!�+�#)�)$�)���&*�()�$#��)���#�3����)�(���0����$���+��($"��'�(�'+�)�$#(��#��'��$""�#��)�$#(2�� ;3 ����'�)��'�%$$'�(�"%!�#����()�$��)���%'$%$(���#�,�'$���!�"�)(�)���%$,�'�)$�)�()���'��)!.��$'�%$)�#)��!���('*%)�$#�$���$##��)�+�).3�<3 ����'�(*!)(��#��$'"�#��)��!3��'��($"�,��)��$*#)�'4�#)*�)�+���#�(*���()�#�������'��$#)�"%$'�'.���(%�'(�!�'�)�(��)�!�'���)��#�("�!!�(��!�3������'��#�!.(�(��$�#$)��#��'�)�����'��)�$#�$����(%�'(�!0�($�,��)��'�)���($*)��'#�%$%*!�)�$#(��'����($*'����$'�)�$(��)$�)���,�()�'�"��#�(%��*!�)�$#3�=3 �$����'�((�7;8��#�)����$#)�-)�$��)����$')��$"�#�����0��)�(�$*!�����%$((��!��)$�"$��!���(%�'(�!�'�)��7*(�#��!$��()���'��'�((�$#�$#�� ����()�#��(�$'�'��'�((�$#�$��%��',�(��'�+�!*�(8��(����*#�)�$#�$��!�#��'���()�#����#������)�)���)�'$��#��).��#��,�)��$'�,�)�$*)���%$)�#)��!�'$�����''��'�>3 �$����'�((�7<80�"�)�$�(�)��)��#��'�"��'�)�$#�'�)�(�$+�'�'���#)���#�'�)�$#(�7�3�3���.�(�((8��$*!��*(���)$�)�()��$'��(.""�)'.��"$#��#$')�0�($*)���#��������#)�'���$#�!�%$%*!�)�$#(3� � Yours truly,