SEXANDTHEC
ITY
:EFFECTSOFIND
IV
IDUALCOND
IT
IONANDHAB
ITAT
URBAN
IZAT
IONON MOUNTA
INCH
ICKADEEREPRODUCT
IVEBEHAV
IOUR
by
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r
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caS
eonBond
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rud
B
.S
c
.Hon
s
.
,Thomp
sonR
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sUn
iv
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r
s
i
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,2015

THES
ISSUBM
ITTEDINPART
IALFULF
ILLMENTOF
THEREQU
IREMENTSFORTHEDEGREEOF
MASTEROFSC
IENCE
IN
NATURALRESOURCESANDENV
IRONMENTALSTUD
IES(B
IOLOGY
)

UN
IVERS
ITYOFNORTHERNBR
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ISHCOLUMB
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rud
,2017

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
ABSTRACT

V
a
r
i
a
t
ioninind
iv
idu
a
lcond
i
t
ionandh
ab
i
t
a
tqu
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l
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lu
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ep
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ego
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er
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ck
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s
.Sp
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ed
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edmoun
t
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ab
i
t
a
tu
rb
an
i
z
a
t
ion
.How
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r
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l
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l
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u
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s
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r
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lh
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i
t
a
ti
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l
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th
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a
r
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z
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t
iono
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emi
s
n
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a
ry
.

i
i

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
PREFACE

Fund
ingfo
rth
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e
s
e
a
r
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a
sp
rov
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i
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cCo
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l
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t
ionP
e
rm
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tno
.22806
.

Th
ed
e
s
ign
,ex
e
cu
t
ion
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ly
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so
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edbyE
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rodu
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sa
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ew
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r
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s
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e
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a
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romth
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r
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l
.

Ch
ap
t
e
r2
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e
rud
,E
.S
.
,O
t
t
e
r
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.A
.
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rg
,T
.M
.
,M
a
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,K
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.D
.&R
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ink
,M
.W
.
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)
.P
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t
t
e
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t
ern
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tyinth
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t
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ck
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e
.E
tho
logy(
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ew
)
.

Ch
ap
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e
r3
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rud
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.
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e
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ay
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rg
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.
,M
a
r
in
i
,K
.L
.D
.&
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ink
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.W
.(2017
)
.E
f
f
e
c
t
so
fp
a
r
en
t
a
lcond
i
t
ionandn
e
s
t
ingh
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t
a
tons
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l
lo
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a
t
ionin
t
h
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t
a
inch
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ck
ad
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e
.–B
eh
aviou
r154
:1101
1121
.

i
i
i

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
TABLEOFCONTENTS

Ab
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A
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v
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Chap
t
e
r1
:	
  G
en
e
ra
lIn
t
rodu
c
t
ion.
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1	
  
1
.1
.	
  U
rb
ane
co
logyandsongb
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rd
s.
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1	
  
1
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3	
  
1
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4	
  
1
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5	
  
1
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6	
  
1
.4
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  S
tudys
i
t
e.
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8	
  
1
.5
.	
  H
ab
i
t
a
tc
l
a
s
s
i
f
i
c
a
t
ion.
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9	
  
1
.6
.	
  Th
e
s
i
sov
e
rv
i
ew.
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11	
  
1
.7
.	
  L
i
t
e
r
a
tu
r
ec
i
t
ed.
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11	
  
1
.8
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  F
igu
r
e
s.
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16	
  
Chap
t
e
r2
:	
  Pa
t
t
e
rn
so
fEx
t
rapa
i
rPa
t
e
rn
i
ty.
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17	
  
2
.1
.	
  Ab
s
t
r
a
c
t.
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17	
  
2
.2
.	
  In
t
rodu
c
t
ion
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17	
  
2
.3
.	
  M
e
thod
s.
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20	
  
2
.3
.1
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  F
i
e
ldm
e
thod
s.
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20	
  
2
.3
.2
.	
  D
awncho
ru
sr
e
co
rd
ing
s.
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21	
  
2
.3
.3
.	
  P
a
r
en
t
a
lc
a
r
e.
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21	
  
2
.3
.4
.	
  Mo
l
e
cu
l
a
rm
e
thod
s.
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22	
  
2
.3
.5
.
	
  P
a
r
en
t
ag
ea
s
s
ignm
en
t.
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23	
  
2
.3
.6
.	
  S
t
a
t
i
s
t
i
c
a
lan
a
ly
s
e
s.
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24	
  
2
.4
.	
  R
e
su
l
t
s.
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26	
  
2
.4
.1
.	
  G
en
e
r
a
lp
a
t
t
e
rn
so
fp
a
r
en
t
ag
e.
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26	
  
2
.4
.2
.
	
  Cond
i
t
ionandex
t
r
ap
a
i
rp
a
t
e
rn
i
ty.
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27	
  
2
.4
.3
.	
  So
c
i
a
lm
a
l
ev
s
.ex
t
r
ap
a
i
rm
a
l
ecomp
a
r
i
son
s.
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27	
  
2
.4
.4
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  H
ab
i
t
a
te
f
f
e
c
t
s.
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27	
  
2
.4
.5
.	
  Ex
t
r
ap
a
i
rp
a
t
e
rn
i
tyandp
a
r
en
t
a
lc
a
r
e.
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28	
  
2
.5
.	
  D
i
s
cu
s
s
ion.
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28	
  
2
.6
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  L
i
t
e
r
a
tu
r
ec
i
t
ed.
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31	
  
2
.7
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  T
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2
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37	
  

iv

E
r
i
c
aS
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e
rud

M
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c
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e
s
i
s (2017
)

Chap
t
e
r3
:	
  E
f
f
e
c
t
so
fPa
r
en
ta
lCond
i
t
ionandU
rban
i
za
t
iononS
exA
l
lo
ca
t
ion.
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40	
  
3
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40	
  
3
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40	
  
3
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42	
  
3
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42	
  
3
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42	
  
3
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42	
  
3
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42	
  
3
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42	
  
3
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43	
  
3
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43	
  
3
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46	
  
3
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t
ionandb
roods
exr
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46
	
  
3
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t
ionandn
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46	
  
3
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eandn
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46	
  
3
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47	
  
3
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  B
roods
exr
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t
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a
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47	
  
3
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47	
  
3
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3
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55
	
  
3
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58	
  
Chap
t
e
r4
:	
  Con
c
lu
s
ion
sandFu
tu
r
eD
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r
e
c
t
ion
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60	
  
4
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tu
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62	
  
4
.2
.	
  Con
c
lu
s
ion
s.
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.
64	
  
4
.3
.	
  L
i
t
e
r
a
tu
r
ec
i
t
ed.
.
.
.
.
.
.
.
.
.
.
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64	
  

v

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
L
ISTOFTABLES

T
ab
l
e2
.1
.A
l
l
e
l
i
cv
a
r
i
a
t
iona
tth
e6m
i
c
ro
s
a
t
e
l
l
i
t
em
a
rk
e
r
s
.S
i
z
er
ang
e
sfo
r
m
i
c
ro
s
a
t
e
l
l
i
t
ea
l
l
e
l
e
sa
r
eg
iv
eninb
a
s
ep
a
i
r
s
,a
longw
i
thth
eto
t
a
lnumb
e
ro
f
un
iqu
ea
l
l
e
l
e
sob
s
e
rv
ed
,andob
s
e
rv
ed(H
ndexp
e
c
t
ed(H
)h
e
t
e
ro
zygo
s
i
t
i
e
s
.
o)a
e

34

T
ab
l
e2
.2
.R
e
su
l
t
so
fg
en
e
r
a
l
i
z
edl
in
e
a
rm
ix
edmod
e
l
sa
sk
inghowm
al
econd
i
t
ion
(
ag
e
,m
a
s
sxt
a
r
su
sr
eg
r
e
s
s
ionr
e
s
idu
a
l
,cho
ru
sch
a
r
a
c
t
e
r
i
s
t
i
c
s
)
,f
em
a
l
e
cond
i
t
ion(
ag
e
,m
a
s
sxt
a
r
su
sr
eg
r
e
s
s
ionr
e
s
idu
a
l
)
,andh
ab
i
t
a
tch
a
r
a
c
t
e
r
i
s
t
i
c
s
(h
ab
i
t
a
tind
ex
,v
eg
e
t
a
t
ionind
ex
)in
f
lu
en
c
eth
ep
r
e
s
en
c
eo
rp
ropo
r
t
iono
fEPOin
n
e
s
t
s
.

35

T
ab
l
e2
.3
.R
e
su
l
t
so
fg
en
e
r
a
l
i
z
edl
in
e
a
rmod
e
l
sa
sk
ingwh
e
th
e
rp
a
r
en
t
a
l
p
rov
i
s
ion
ingi
sad
ju
s
t
edinr
e
spon
s
etoth
ep
r
e
s
en
c
eo
rp
ropo
r
t
iono
fEPOin
n
e
s
t
s
.

36

T
ab
l
e3
.1
.Summ
a
ryo
fmoun
t
a
inch
i
ck
ad
e
es
exr
a
t
iod
a
t
afo
rth
e2014
–2016
b
r
e
ed
ings
e
a
son
s
.W
i
l
coxons
ign
ed
r
ankt
e
s
t
sw
e
r
ep
e
r
fo
rm
edtod
e
t
e
rm
in
ei
f
popu
l
a
t
iono
rb
roods
exr
a
t
io
sd
ev
i
a
t
edf
romun
i
ty
.As
ing
l
ec
a
s
eo
fr
en
e
s
t
ing
w
a
sex
c
lud
edf
rom2016an
a
ly
s
e
s
.

55

T
ab
l
e3
.2
.R
e
su
l
t
so
fg
en
e
r
a
l
i
z
edl
in
e
a
rm
ix
edmod
e
l
sex
am
in
ingth
ein
f
lu
en
c
eo
f
so
c
i
a
lm
a
l
eandg
en
e
t
i
cf
a
th
e
rcond
i
t
iononn
e
s
t
l
ings
ex
.

56

T
ab
l
e3
.3
.R
e
su
l
t
so
fg
en
e
r
a
l
i
z
edl
in
e
a
rm
ix
edmod
e
l
sex
am
in
ingth
ein
f
lu
en
c
eo
f
f
em
a
l
econd
i
t
iononn
e
s
t
l
ings
ex
.

57

v
i

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
L
ISTOFF
IGURES

F
igu
r
e1
.1
.Lo
c
a
t
ion
so
fa
l
lmoun
t
a
inch
i
ck
ad
e
en
e
s
tbox
e
sinK
am
loop
s
,BC
,
C
an
ad
a(boxA
)
.N
e
s
tbox
e
sw
e
r
ed
i
s
t
r
ibu
t
edth
roughou
tsou
thK
am
loop
sona
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ab
i
t
a
tg
r
ad
i
en
tf
romru
r
a
ltou
rb
an
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c
a
t
ion
sin
c
lud
edth
econ
i
f
e
rfo
r
e
s
t
so
f
K
enn
aC
a
r
tw
r
igh
tP
a
rk(
i
.e
.
,ru
r
a
l
;r
edpo
in
t
s
,boxB
)
,subu
rb
an
n
e
ighbou
rhood
s(g
r
e
enpo
in
t
s
;boxA
)
,andth
eThomp
sonR
iv
e
r
sUn
iv
e
r
s
i
ty
C
ampu
s(
i
.
e
.
,u
rb
an
;b
lu
epo
in
t
s
,boxC)
.

16

F
igu
r
e2
.1
.(A
)P
e
r
c
en
t
ag
eo
fn
e
s
t
scon
t
a
in
inga
tl
e
a
s
ton
eex
t
r
ap
a
i
ro
f
f
sp
r
ing
(EPO
)anda
l
lw
i
th
inp
a
i
ro
f
f
sp
r
ing(WPO
)
;(B
)P
e
r
c
en
t
ag
eo
fo
f
f
sp
r
ingth
a
t
w
e
r
eex
t
r
ap
a
i
randw
i
th
inp
a
i
r
.Numb
e
r
sw
i
th
inb
a
r
sr
ep
r
e
s
en
t
sto
t
a
lnumb
e
r
o
f(A
)n
e
s
t
sand(B
)o
f
f
sp
r
ings
amp
l
edine
a
chs
tudyy
e
a
r
.

37

F
igu
r
e2
.2.Th
ep
ropo
r
t
iono
fEPOinan
e
s
tw
a
sno
ti
n
f
lu
en
c
edbye
i
th
e
r(A
)th
e
h
ab
i
t
a
tind
exo
r(B
)v
eg
e
t
a
t
ionind
exo
fth
en
e
s
t
ingh
ab
i
t
a
t
.Th
e
s
eind
i
c
e
sw
e
r
e
d
e
r
iv
edtod
e
s
c
r
ib
e(A
)th
ed
eg
r
e
eo
fh
ab
i
t
a
tu
rb
an
i
z
a
t
ionandov
e
r
a
l
lqu
a
l
i
ty
,
and(B
)v
eg
e
t
a
t
ioncompo
s
i
t
ion
,c
anopycov
e
randfoodav
a
i
l
ab
i
l
i
ty
.

38

F
igu
r
e2
.3.N
e
s
t
scon
t
a
in
ingag
r
e
a
t
e
rp
ropo
r
t
iono
fEPOw
e
r
ep
rov
i
s
ion
eda
th
igh
e
r
r
a
t
e
s
.Th
i
sr
e
l
a
t
ion
sh
ipw
a
ss
ign
i
f
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c
an
twh
enbo
thn
e
s
t
sw
i
thandw
i
thou
tEPO
w
e
r
econ
s
id
e
r
ed(p=0
.007
;b
l
a
ckl
in
e
)
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em
a
in
eds
ign
i
f
i
c
an
twh
enw
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p=0
.006
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r
eyl
in
e
)
.
i
so
l
a
t
edn
e
s
t
scon
t
a
in
ingEPO(

39

F
igu
r
e3
.1
.D
i
s
t
r
ibu
t
iono
fb
roods
exr
a
t
io
sfo
rth
e20142016b
r
e
ed
ings
e
a
son
s
.
B
roods
exr
a
t
io
sr
ang
edf
rom0
.17(
a
lmo
s
tex
c
lu
s
iv
e
lyf
em
a
l
e
)to1
.0
(
ex
c
lu
s
iv
e
lym
a
l
e
)
.

58

F
igu
r
e3
.2
.H
ab
i
t
a
th
adane
f
f
e
c
tonn
e
s
t
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ings
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c
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r
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l
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s
)
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tno
tru
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enc
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l
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ab
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t
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t
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low
e
rh
ab
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t
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ind
exs
co
r
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s
)
.

59

v
i
i

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
ACKNOWLEDGEMENTS

Iwou
ldl
ik
etoa
cknow
l
edg
eth
i
sr
e
s
e
a
r
chw
a
scondu
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edonth
et
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ad
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a
lt
e
r
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to
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fth
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em
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ép
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i
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iontocondu
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tr
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a
r
chinK
enn
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ingm
etomon
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v
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i
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E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)
CHAPTER1
:GENERALINTRODUCT
ION

1
.1
. U
rbane
co
logyandsongb
i
rd
s
f
Ou
rp
l
an
e
ti
sr
ap
id
lyb
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com
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rb
an
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z
ed
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e
tw
e
en1970and2000
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e
a
r
ly60
,000km2o
l
andw
a
sconv
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r
t
edtou
rb
anu
s
e
,andth
i
sa
r
e
ai
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j
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c
t
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c
r
e
a
s
ebyov
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r1
.5m
i
l
l
ion
2
km
byth
ey
e
a
r2030(S
e
toe
ta
l
.2011
)
.Thou
ghd
ev
e
lop
ingcoun
t
r
i
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sa
r
eexp
e
r
i
en
c
ingth
e

g
r
e
a
t
e
s
tin
c
r
e
a
s
einu
rb
an
i
z
a
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ion
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rb
anl
anda
r
e
acon
t
inu
e
stor
i
s
einNo
r
thAm
e
r
i
c
aand
Eu
rop
e(S
e
toe
ta
l
.2011
)
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l
r
e
ady
u
rb
an
i
z
ed
,d
ev
e
lop
edcoun
t
r
i
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s
,th
epopu
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a
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ion
sa
r
e
in
c
r
e
a
s
ing
lymov
ingou
to
fu
rb
anc
en
t
r
e
sandr
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s
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t
t
l
inginsubu
rb
ana
r
e
a
s
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rb
an
co
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i
z
a
t
ioni
sch
a
r
a
c
t
e
r
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edbysp
r
aw
l
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,low
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d
en
s
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tys
e
t
t
l
em
en
tth
anu
rb
an
,and
con
s
equ
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t
ly
,c
i
t
i
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sind
ev
e
lop
edcoun
t
r
i
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sa
r
eg
row
ingf
a
s
t
e
rina
r
e
ath
aninpopu
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a
t
ion
(M
a
r
z
lu
f
fe
ta
l
.2001
)
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ssu
ch
,i
ti
simpo
r
t
an
tw
efu
r
th
e
rou
rund
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r
s
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l
l
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re
ta
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l
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r
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s
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1

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r
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rud

M
.S
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s
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s (2017
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l
l
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re
ta
l
.2012
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l
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r
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s
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).
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ly
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l
a
i
r1996
;S
e
r
e
s
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r
2015
).U
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e
r
e
s
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amp
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e
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u
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cha
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em
ig
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a
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ryin
s
e
c
t
ivo
r
e
s(B
l
a
i
r1996
)
.
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esp
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c
i
e
s
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r
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r
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e
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s
s
e
rdom
e
s
t
i
cu
s)(B
l
a
i
r1996
)
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e
s
esp
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c
i
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s
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r
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.
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i
rdf
e
ed
e
r
s
,d
i
spo
s
edfood
)(B
l
a
i
r1996
;S
e
r
e
s
sandL
ik
e
r2015
)
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ef
in
a
l
c
a
t
ego
ry
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rb
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s
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i
s
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s
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l
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s
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e
r
s
)(B
l
a
i
r
1996
;S
e
r
e
s
sandL
ik
e
r2015
)
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rb
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ap
to
r
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r
ecommon
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n
e
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i
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e
r
e
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t
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e
c
i
l
eru
f
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s
c
en
s)(B
l
a
i
r1996
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.
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etoi
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t
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2

E
r
i
c
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.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

d
e
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i
s
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s
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r
ap
a
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rcopu
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a
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s(EPC
s
)(G
r
i
f
f
i
the
ta
l
.2002
)
.
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r
s
e
ly
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fu
rb
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z
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t
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c
r
e
a
s
e
se
i
th
e
rd
en
s
i
ty(M
a
r
z
lu
f
f1997
)o
raf
em
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l
e
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a
s
s
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s
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a
l
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l
s(L
aZ
e
r
t
ee
ta
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.2015
)
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a
t
ing
sm
ayb
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c
ed
.F
em
a
l
e
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ly
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ik
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i
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s(S
l
abb
ekoo
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l
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t(L
aZ
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r
t
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ta
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.2017
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t
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s(T
r
iv
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r
sand W
i
l
l
a
rd1973
)
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sonandBu
r
l
ey1998
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lr
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iv
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tpu
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l
e
s(T
r
iv
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r
s1972
)
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ths
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sa
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ay
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a
t
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ax
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f
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r
iv
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r
s
3

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

1972
).Todoso
,f
em
a
l
e
sm
ay
,fo
rex
amp
l
e
,
eng
ag
einEPC
sw
i
thm
a
l
e
so
fh
ighqu
a
l
i
ty
(G
r
i
f
f
i
the
ta
l
.2002
;W
e
s
tn
e
a
tandS
t
ew
a
r
t2003
)o
rb
i
a
so
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f
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l(T
r
iv
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r
sand W
i
l
l
a
rd1973
;A
lon
so
A
lv
a
r
e
z2006
).Ex
t
r
ap
a
i
rp
a
t
e
rn
i
ty(
EPP)andb
roods
exr
a
t
iob
i
a
sa
r
eju
s
ttwoo
fth
em
anyw
ay
sf
em
a
l
e
sm
ay
in
c
r
e
a
s
er
ep
rodu
c
t
iv
esu
c
c
e
s
s
,andth
eyw
i
l
lb
eth
efo
cu
so
fth
i
sth
e
s
i
s
.

1
.2
.1
.E
x
t
rapa
i
rpa
t
e
rn
i
t
y
Th
ev
a
s
tm
a
jo
r
i
tyo
fsongb
i
rdsp
e
c
i
e
sa
r
eso
c
i
a
l
lymonog
amou
s
,w
i
thas
ing
l
em
a
l
eand
f
em
a
l
ep
a
i
r
ingandc
a
r
ingfo
ro
f
f
sp
r
ing(W
e
s
tn
e
a
tandS
t
ew
a
r
t2003
)
.T
r
iv
e
r
s(1972
)
sugg
e
s
t
edth
a
tm
a
l
e
sinso
c
i
a
l
lymonog
amou
ssp
e
c
i
e
sm
ayp
a
r
t
i
c
ip
a
t
einam
ix
edm
a
t
ing
s
t
r
a
t
egyo
fso
c
i
a
lmonog
amywh
i
l
ein
c
r
e
a
s
ingr
ep
rodu
c
t
iv
esu
c
c
e
s
sth
roughEPC
sw
i
th
n
e
ighbou
r
ingf
em
a
l
e
s
,s
i
r
ingo
f
f
sp
r
ingbu
tw
i
thho
ld
ingp
a
r
en
t
a
lc
a
r
e
.L
ik
ew
i
s
e
,so
c
i
a
l
ly
monog
amou
sf
em
a
l
e
sm
ayin
c
r
e
a
s
er
ep
rodu
c
t
iv
esu
c
c
e
s
sbyeng
ag
inginEPC
s(T
r
iv
e
r
s
1972
)
.Th
eadv
en
to
fmo
l
e
cu
l
a
rtoo
l
sfo
rp
a
t
e
rn
i
tyan
a
ly
s
i
sr
ev
e
a
l
edth
e
s
ep
r
ed
i
c
t
eds
t
r
a
t
eg
i
e
s
tob
ew
id
e
sp
r
e
ad(B
i
rkh
e
adand Mø
l
l
e
r1992
)
,w
i
th
a
tl
e
a
s
t70%o
fso
c
i
a
l
lymonog
amou
s
songb
i
rdsp
e
c
i
e
scon
f
i
rm
edtob
eg
en
e
t
i
c
a
l
l
yp
rom
i
s
cuou
s(W
e
s
tn
e
a
tandS
t
ew
a
r
t2003
)
.
Th
e
r
ea
r
es
ev
e
r
a
lhypo
th
e
s
e
sa
stowhyaf
em
a
l
em
aychoo
s
etoeng
ag
einEPC
s
in
c
lud
ing
:toen
su
r
ef
e
r
t
i
l
i
z
a
t
iono
fh
e
regg
s
;toin
c
r
e
a
s
eth
eg
en
e
t
i
cd
iv
e
r
s
i
tyo
rg
en
e
t
i
c
qu
a
l
i
tyo
fh
e
ro
f
f
sp
r
ing
;o
r
,tor
e
c
e
iv
ed
i
r
e
c
tb
en
e
f
i
t
s
,l
ik
ed
e
f
en
s
eo
rr
e
sou
r
c
e
s
,f
romth
eEP
m
a
l
e(G
r
i
f
f
i
the
ta
l
.2002
)
.On
ehypo
th
e
s
i
sth
a
th
a
sg
a
rn
e
r
eds
ign
i
f
i
c
an
ta
t
t
en
t
ioni
sth
egood
g
en
e
shypo
th
e
s
i
s(B
i
rkh
e
adand Mø
l
l
e
r1992
;K
emp
en
a
e
r
sandDhond
t1993
)
.Th
egood
g
en
e
shypo
th
e
s
i
ss
t
a
t
e
sth
a
tf
em
a
l
ess
e
ekEPC
sw
i
thm
a
l
e
so
fg
r
e
a
t
e
rg
en
e
ti
cqu
a
l
i
tyth
an
th
e
i
rso
c
i
a
lm
a
t
eino
rd
e
rtoob
t
a
inmo
r
ef
avou
r
a
b
l
eg
en
e
sfo
rth
e
i
ro
f
f
sp
r
ing(K
emp
en
a
e
r
s
andDhond
t1993
;G
r
i
f
f
i
the
ta
l
.2002
)
.Und
e
rth
i
shypo
th
e
s
i
s
,f
em
a
l
e
sa
s
s
e
s
sm
a
l
econd
i
t
ion
4

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

th
roughcond
i
t
ion
d
ep
end
en
tph
eno
typ
i
ccu
e
s(K
emp
en
a
e
r
sandDh
ond
t1993
;G
r
i
f
f
i
the
ta
l
.
2002
).Inadd
i
t
ion
,h
ab
i
t
a
tqu
a
l
i
tyi
sa
l
socommon
lyhypo
th
e
s
i
z
edtoin
f
lu
en
c
er
a
t
e
so
fEPP
.
Inlow
qu
a
l
i
tyh
ab
i
t
a
t
sw
i
thl
im
i
t
edfoodav
a
i
l
ab
i
l
i
ty
,EPPi
sp
r
ed
i
c
t
edtob
emo
r
ep
r
ev
a
l
en
t
,
a
sf
em
a
l
e
sm
ayeng
ag
einEPC
sw
i
thm
a
le
sonh
igh
e
r
qu
a
l
i
tyt
e
r
r
i
to
r
i
e
sino
rd
e
rtog
a
in
in
c
r
e
a
s
eda
c
c
e
s
stofood(G
r
i
f
f
i
the
ta
l
.2002
)
.Th
ep
r
e
s
en
c
eo
fEPP
insongb
i
rd
si
s
common
lya
t
t
r
ibu
t
edtod
i
f
f
e
r
en
c
e
sinm
a
t
econd
i
t
iono
rqu
a
l
i
ty(
e
.g
.
,K
emp
en
a
e
r
se
ta
l
.
1992
,1997
;O
t
t
e
re
ta
l
.1998
)o
rt
e
r
r
i
to
ryqu
a
l
i
ty(
e
.g
.
,Rub
en
s
t
e
in2007
)
.

1
.2
.2
.S
e
xa
l
lo
ca
t
ion
S
exa
l
lo
c
a
t
ionth
eo
ryp
r
ed
i
c
t
sf
em
a
l
e
sshou
ldb
i
a
sth
es
exr
a
t
ioo
fth
e
i
ro
f
f
sp
r
inginr
e
spon
s
e
tod
i
f
f
e
r
en
c
e
sinth
er
ep
rodu
c
t
iv
ev
a
lu
eo
fson
sandd
augh
t
e
r
s(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;
Ch
a
rnov1982
).B
e
c
au
s
em
a
l
e
styp
i
c
a
l
lyexp
e
r
i
en
c
eg
r
e
a
t
e
rv
a
r
i
an
c
einr
ep
rodu
c
t
iv
esu
c
c
e
s
s
th
anf
em
a
l
e
s
,ah
igh
qu
a
l
i
tysonm
ayh
av
eth
epo
t
en
t
i
a
ltop
rodu
c
emo
r
eo
f
f
sp
r
ingandh
av
e
h
igh
e
rr
ep
rodu
c
t
iv
esu
c
c
e
s
sth
anad
augh
t
e
ro
fth
es
am
equ
a
l
i
ty
,wh
i
l
eth
eoppo
s
i
t
em
ayb
e
t
ru
efo
rlow
qu
a
l
i
tyson
sandd
augh
t
e
r
s(T
r
iv
e
r
sand W
i
l
l
a
rd1973
)
.O
f
f
sp
r
ingr
ep
rodu
c
t
iv
e
v
a
lu
em
ayv
a
rya
sar
e
su
l
to
fbo
thin
t
r
in
s
i
cf
a
c
to
r
s(
e
.g
.
,th
ef
em
a
l
e
’
sowncond
i
t
ion
)and
ex
t
r
in
s
i
cf
a
c
to
r
s(
e
.g
.
,m
a
t
econd
i
t
ionandh
ab
i
t
a
tqu
a
l
i
ty
)(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;Bu
r
l
ey
1981
).T
h
e
r
e
fo
r
e
,f
em
a
l
e
sshou
ldb
i
a
sb
roods
exr
a
t
io
sinr
e
spon
s
etosu
cha
t
t
r
ibu
t
e
s
.In
n
a
tu
r
e
,b
i
a
s
eds
exa
l
lo
c
a
t
ioninr
e
spon
s
etop
a
r
en
t
a
lcond
i
t
ionandh
ab
i
t
a
tqu
a
l
i
tyh
a
sb
e
en
do
cum
en
t
edinm
anyv
e
r
t
eb
r
a
t
e
s
,in
c
lud
ingnum
e
rou
sb
i
rdsp
e
c
i
e
s(
r
ev
i
ew
edinA
lon
so
A
lv
a
r
e
z2006
).
Th
eT
r
iv
e
r
sand W
i
l
l
a
rd(1973
)hypo
th
e
s
i
so
fs
exa
l
lo
c
a
t
ionp
r
ed
i
c
t
sf
em
a
l
e
singood
cond
i
t
ionshou
ldp
rodu
c
em
a
l
e
b
i
a
s
edb
rood
s
,a
s
s
um
ingo
f
f
spr
ingcond
i
t
ionandsu
rv
iv
a
la
r
e
co
r
r
e
l
a
t
edw
i
thm
a
t
e
rn
a
lcond
i
t
ion
.Th
eCh
a
rnovmod
e
l(1982
)p
r
ed
i
c
t
sfem
a
l
e
sshou
lda
l
t
e
r
5

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

s
exa
l
lo
c
a
t
ionb
a
s
edonp
a
t
e
rn
a
la
t
t
r
ibu
t
e
sth
a
tm
ayb
en
e
f
i
ton
es
exmo
r
eth
anth
eo
th
e
r
.Th
e
r
e
l
a
t
ion
sh
ipb
e
tw
e
enm
a
t
e
rn
a
lcond
i
t
ionandb
roods
exr
a
t
ioh
a
sb
e
enw
e
l
ls
tud
i
edand
suppo
r
t
edins
ev
e
r
a
lav
i
ansp
e
c
i
e
s(N
ag
e
re
ta
l
.1999
; Wh
i
t
t
ingh
amandDunn2000
;A
lon
so
A
lv
a
r
e
zandV
e
l
ando2003
;P
ik
eandP
e
t
r
i
e2005
).S
tud
i
e
sonsongb
i
rdsp
e
c
i
e
sh
av
e
common
lycon
s
id
e
r
edth
ein
f
lu
en
c
eo
fm
a
l
econd
i
t
ionandh
ab
i
t
a
tqu
a
l
i
tyono
f
f
sp
r
ings
ex
.
Fo
rex
amp
l
e
,m
a
l
emoun
t
a
inb
lu
eb
i
rd
s(S
ia
l
iacu
r
ru
co
id
e
s)d
i
sp
l
ayfu
l
l
bodyUVb
lu
e
p
lum
ag
e
,th
eco
lou
ro
fwh
i
chi
sd
ep
end
e
n
tonanind
iv
idu
a
l
’
snu
t
r
i
t
ion
a
ls
t
a
t
ea
tth
et
im
eo
f
mou
l
t(
i
.
e
.
,cond
i
t
iond
ep
end
en
t
)(K
ey
s
e
randH
i
l
l1999
;S
i
e
f
f
e
rm
anandH
i
l
l2007
;Doy
l
e
andS
i
e
f
f
e
rm
an2014
)
.F
em
a
l
emoun
t
a
in
b
lu
eb
i
rd
sp
a
i
r
edw
i
thmo
r
eb
r
igh
t
lyco
lou
r
edm
a
l
e
s
p
rodu
c
em
a
l
eb
i
a
s
edb
rood
s
,sugg
e
s
t
ingf
em
a
l
e
sp
e
r
c
e
iv
eandr
e
spondtov
a
r
i
a
t
ioninm
a
l
e
co
lou
r
a
t
ion(
i
.
e
.
,cond
i
t
ion
)(Bond
e
rude
ta
l
.2016
)
.
Th
ephy
s
io
log
i
c
a
lm
e
ch
an
i
sm
sa
l
low
ingf
em
a
l
e
stop
rodu
c
eb
i
a
s
edb
roods
exr
a
t
io
sa
t
l
ay
inga
r
epoo
r
lyund
e
r
s
tood
,how
ev
e
r
,th
emo
s
tp
l
au
s
ib
l
eth
eo
r
i
e
ssugg
e
s
tf
em
a
l
eho
rmon
a
l
con
t
ro
l(P
ik
eandP
e
t
r
i
e2003
;N
av
a
r
a2013
)
.T
e
s
to
s
t
e
ron
e
,fo
rex
amp
l
e
,i
sp
r
e
s
en
ting
r
e
a
t
e
r
amoun
t
sinth
eyo
lk
so
fp
e
a
fow
l(
Pa
voc
r
i
s
ta
tu
s)egg
scon
t
a
in
ingm
a
l
ech
i
ck
s
,andh
a
sb
e
en
sugg
e
s
t
edt
op
romo
t
eth
ed
ev
e
lopm
en
to
fm
a
l
eegg
sf
romth
ef
i
r
s
tm
e
io
t
i
cd
iv
i
s
ion(P
e
t
r
i
ee
t
a
l
.2001
)
.F
em
a
l
eho
rmon
el
ev
e
l
sch
ang
einr
e
spon
s
etom
a
t
ea
t
t
r
a
c
t
iv
en
e
s
s
,s
e
l
f
cond
i
t
ion
anda
t
t
r
a
c
t
iv
en
e
s
s
,andr
e
sou
r
c
eav
a
i
l
ab
i
l
i
ty
,thu
s
,th
e
s
eex
t
e
rn
a
lf
a
c
to
r
sm
ayb
er
e
f
l
e
c
t
edin
ov
a
r
i
anandyo
lkho
rmon
el
ev
e
l
s
,r
e
su
l
t
ingins
exsp
e
c
i
f
i
cfo
l
l
i
c
l
eo
remb
ryod
ev
e
lopm
en
t
(P
ik
eandP
e
t
r
i
e2003
)
.

1
.3
.S
tudysp
e
c
i
e
s
Moun
t
a
inch
i
ck
ad
e
e
sa
r
esm
a
l
l
,non
m
ig
r
a
to
rysongb
i
rd
scommontoth
emon
t
an
econ
i
f
e
rou
s
fo
r
e
s
t
so
fw
e
s
t
e
rnNo
r
thAm
e
r
i
c
a(M
cC
a
l
lume
ta
l
.1999
)
.Th
i
sh
igh
lyad
ap
t
ab
l
esp
e
c
i
e
sa
l
so
6

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

common
lyco
lon
i
z
e
ssubu
rb
anh
ab
i
t
a
t
. Moun
t
a
inch
i
ck
ad
e
e
sa
r
es
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sgov
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in
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c
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i
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r
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ch
i
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s(M
cC
a
l
lume
ta
l
.1999
;R
a
t
c
l
i
f
f
ee
ta
l
.2007
)
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l
thoughmu
cho
fou
r
und
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cC
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l
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ta
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.1999
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l
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l
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r
av
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ta
l
.2012
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r
te
ta
l
.2007
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a
l
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t(O
t
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re
ta
l
.1997
)
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l
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ta
l
.2007
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t
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t
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a
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c
l
i
f
f
e1996
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am
s
ay
a
r
tn
e
r
s(O
t
t
e
re
ta
l
.1994
,1998
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.In
c
r
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s
eda
c
c
e
s
stor
e
sou
r
c
e
styp
i
c
a
l
ly
e
ta
l
.2000
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l
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igh
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rv
ivo
r
sh
ip
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r
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t
t
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re
ta
l
.1999
;S
chub
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r
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ta
l
.2007
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edom
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rd
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ighr
ank
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chub
e
r
te
ta
l
.2008
)r
e
l
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etoyoung
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a
l
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sa
l
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s
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e
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sm
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equ
a
l
i
ty(O
t
t
e
r
e
ta
l
.1997
;M
enn
i
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le
ta
l
.2003
)
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l
a
ck
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app
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i
ck
ad
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l
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e
m
a
l
e
s
)(O
t
t
e
re
ta
l
.1997
;G
r
av
ae
ta
l
.2009
)
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em
en
t
a
lf
e
ed
ingexp
e
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im
en
t
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r
av
ae
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a
l
.(2009
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7

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r
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rud

M
.S
c
.Th
e
s
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s (2017
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subo
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igh
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l
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r
ibu
t
ingtov
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r
i
a
t
ioninsongou
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t(G
r
av
ae
ta
l
.2009
)
.
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s
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f
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r
en
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einsongou
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tb
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e
endom
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r
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i
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a
r
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ta
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.2017
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t
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iv
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r
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t
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cC
a
l
lume
ta
l
.1999
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a
th
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iduou
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fo
r
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t
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a
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a
r
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l
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a
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e
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1
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1
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. L
i
t
e
ra
tu
r
ec
i
t
ed
A
lon
soA
lv
a
r
e
z
,C
.(2006
)
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an
ipu
l
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t
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r
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a
rys
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:anupd
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Pou
l
t
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io
l
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ev
.17
:120
.
A
lon
soA
lv
a
r
e
z
,C
.&V
e
l
ando
,A
.(2003
)
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em
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l
ebodycond
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t
ionandb
roods
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a
t
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y
e
l
lowl
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l
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ru
sca
ch
innan
s.— Ib
i
s145
:220
226
.
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.
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e
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.
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eb
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,M
.
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eu
r
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romegg
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:n
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a
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.— J
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i
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r
c
einpu
tch
ang
e
spopu
l
a
t
iondyn
am
i
c
so
fc
i
tys
l
i
ck
e
rb
i
rd
s
.— O
iko
s106
:622626
.

14

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

S
i
e
f
f
e
rm
an
,L
.&H
i
l
l
,G
.E
.(2007
)
.Th
ee
f
f
e
c
to
fr
e
a
r
ingenv
i
ronm
en
tonb
lu
es
t
ru
c
tu
r
a
l
co
lo
r
a
t
iono
fe
a
s
t
e
rnb
lu
eb
i
rd
s(
S
ia
l
ias
ia
l
i
s
)
.— B
eh
av
.E
co
l
.So
c
iob
io
l
.61
:18391846
.
S
l
abb
ekoo
rn
,H
.(2013
)
.Song
so
fth
ec
i
ty
:no
i
s
ed
ep
end
en
tsp
e
c
t
r
a
lp
l
a
s
t
i
c
i
tyinth
ea
cou
s
t
i
c
ph
eno
typ
eo
fu
rb
anb
i
rd
s
.— An
im
.B
eh
av
.85
:10891099
.
Sm
i
th
,S
.M
.(1976
)
.E
co
log
i
c
a
la
sp
e
c
t
so
fdom
in
an
c
eh
i
e
r
a
r
ch
i
e
sinb
l
a
ckc
app
edch
i
ck
ad
e
e
s
.
— Th
eAuk93
:95107
.
Sou
thwood
,T
.R
.E
.(1961
)
.Th
enumb
e
ro
fsp
e
c
i
e
so
fin
s
e
c
t
sa
s
so
c
i
a
t
edw
i
thv
a
r
iou
st
r
e
e
s
.—
J
.An
im
.E
co
l
.30
:1
8
.
T
r
iv
e
r
s
,R
.L
.(1972
)
.P
a
r
en
t
a
linv
e
s
tm
en
tands
exu
a
ls
e
l
e
c
t
ion
.— In
:S
exu
a
ls
e
l
e
c
t
ionand
th
ed
e
s
c
en
to
fm
an(C
ampb
e
l
l
,B
.ed
.
)
.A
ld
in
ePub
l
i
sh
ing
,Ch
i
c
ago
,p
.136179
.
T
r
iv
e
r
s
,R
.L
.& W
i
l
l
a
rd
,D
.E
.(1973
)
.N
a
tu
r
a
ls
e
l
e
c
t
iono
fp
a
r
en
t
a
lab
i
l
i
tytov
a
ryth
es
ex
r
a
t
ioo
fo
f
f
sp
r
ing
.— S
c
i
en
c
e179
:9092
.
W
e
s
tn
e
a
t
,D
.F
.&S
t
ew
a
r
t
,I
.R
.K
.(2003
)
.Ex
t
r
ap
a
i
rp
a
t
e
rn
i
tyinb
i
rd
s:c
au
s
e
s
,co
r
r
e
l
a
t
e
s
,and
con
f
l
i
c
t
.— Annu
.R
ev
.E
co
l
.Evo
l
.S
.34
:365396
.
Wh
i
t
t
ingh
am
,L
.A
.&Dunn
,P
.O
.(2000
)
.O
f
f
sp
r
ings
exr
a
t
io
sint
r
e
esw
a
l
low
s
:f
em
a
l
e
sin
b
e
t
t
e
rcond
i
t
ionp
rodu
c
emo
r
eson
s
.— Mo
l
.E
co
l
.9
:11231129
.

15

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s(2017
)

1
.8
. F
igu
r
e
s

F
igu
r
e1
.1
.Lo
c
a
t
ion
so
fa
l
lmoun
t
a
inch
i
ck
ad
e
en
e
s
tbox
e
sinK
am
loop
s
,BC
,C
an
ad
a(box
A
)
.N
e
s
tbox
e
sw
e
r
ed
i
s
t
r
ibu
t
edth
roughou
tsou
thK
am
loop
sonah
ab
i
t
a
tg
r
ad
i
en
tf
romru
r
a
l
tou
rb
an
.Lo
c
a
t
ion
sin
c
lud
edth
econ
i
f
e
rfo
r
e
s
t
so
fK
enn
aC
a
r
tw
r
igh
tP
a
rk(
i
.
e
.
,ru
r
a
l
;r
ed
po
in
t
s
,boxB
)
,subu
rb
ann
e
ighbou
rhood
s(g
r
e
enpo
in
t
s
;boxA
)
,andth
eThomp
sonR
iv
e
r
s
Un
iv
e
r
s
i
tyC
ampu
s(
i
.
e
.
,u
rb
an
;b
lu
epo
in
t
s
,boxC
)
.

16

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s
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s (2017
)

CHAPTER2
:PATTERNSOFEXTRAPA
IRPATERN
ITY
2
.1
. Ab
s
t
ra
c
t
Ex
t
r
ap
a
i
rp
a
t
e
rn
i
tyi
scommoninP
a
r
id
a
esp
e
c
i
e
sando
f
t
ena
t
t
r
ibu
t
edtoth
egoodg
en
e
s
hypo
th
e
s
i
s
.Ino
th
e
rsongb
i
rdsp
e
c
i
e
s
,h
ab
i
t
a
tqu
a
l
i
tyin
f
lu
en
c
e
sEPPr
a
t
e
s
.H
e
r
e
,w
ea
skhow
ind
iv
idu
a
lcond
i
t
ionandh
ab
i
t
a
tu
rb
an
i
z
a
t
ionin
f
lu
en
c
er
a
t
e
so
fEPPinmoun
t
a
inch
i
ck
ad
e
e
s
.
Ov
e
rth
r
e
eb
r
e
ed
ings
e
a
son
s
,wemo
n
i
to
r
edmoun
t
a
inch
i
ck
ad
e
en
e
s
t
sinu
rb
anandru
r
a
l
h
ab
i
t
a
t
,andd
e
t
e
rm
in
edp
a
r
en
t
ag
ebyg
eno
typ
ingind
iv
idu
a
l
sa
t6m
i
c
ro
s
a
t
e
l
l
i
t
elo
c
i
.EPPi
s
commoninmoun
t
a
inch
i
ck
ad
e
e
s
,w
i
thex
t
r
a
p
a
i
ro
f
f
sp
r
ing(EPO
)p
r
e
s
en
tin44
.4%o
fn
e
s
t
s
anda
c
coun
t
ingfo
r17
.
8%o
fo
f
f
sp
r
ing
.W
efoundsuppo
r
tfo
rth
egoodg
en
e
shypo
th
e
s
i
s
,w
i
th
o
ld
e
rm
a
l
e
st
end
ingtor
a
i
s
ef
ew
e
rEPO
,andf
em
a
l
e
st
end
ingtoeng
ag
einEPC
sw
i
tho
ld
e
r
m
a
l
e
s.P
a
r
en
t
a
lp
rov
i
s
ion
inge
f
fo
r
tw
a
s
g
r
e
a
t
e
rinb
rood
scon
t
a
in
ingag
r
e
a
t
e
rp
ropo
r
t
iono
f
EPO
.W
ed
idno
tf
inda s
ign
i
f
i
c
an
te
f
f
e
c
to
fh
ab
i
t
a
tonEPPr
a
t
e
s
,sugg
e
s
t
ingth
eimp
a
c
t
so
f
u
rb
an
i
z
a
t
iononmoun
t
a
inch
i
ck
ad
e
e
sm
ayb
emo
r
ecomp
l
exth
ano
r
ig
in
a
l
lyp
r
ed
i
c
t
ed.

2
.2
. In
t
rodu
c
t
ion
Inb
i
rd
s
,so
c
i
a
lmonog
amyi
sw
id
e
sp
r
e
ad
,bu
ti
so
f
t
encoup
l
edw
i
tham
ix
ed
m
a
t
ings
t
r
a
t
egy
th
a
tin
c
lud
e
sEPP
.Th
e
r
ea
r
es
ev
e
r
a
lhypo
th
e
s
e
sa
stowhyaf
em
a
l
em
aychoo
s
etoeng
ag
ein
EPC
sin
c
lud
ing
:toi
n
su
r
ef
e
r
t
i
l
i
z
a
t
iono
fh
e
regg
s
;in
c
r
e
a
s
eth
eg
en
e
t
i
cd
iv
e
r
s
i
tyo
fh
e
r
o
f
f
sp
r
ing
;o
r,tor
e
c
e
iv
ed
i
r
e
c
tb
en
e
f
i
t
s
,l
ik
ed
e
f
en
s
eo
rr
e
sou
r
c
e
s
,f
romth
eEPm
a
l
e(G
r
i
f
f
i
th
e
ta
l
.2002
)
.Ex
t
r
ap
a
i
rp
a
t
e
rn
i
tyi
scommoninth
eP
a
r
id
a
ef
am
i
ly(
ch
i
ck
ad
e
e
sandt
i
tm
i
c
e
)
,
w
i
thex
t
r
ap
a
i
ro
f
f
sp
r
ing(EPO
)p
r
e
s
en
tin30
75%o
fn
e
s
t
s
,anda
c
coun
t
ingfo
r725%o
f
o
f
f
sp
r
ing(G
r
i
f
f
i
the
ta
l
.2002
)
.AmongP
a
r
id
s
,EPPi
so
f
t
enexp
l
a
in
edbyth
egoodg
en
e
s
hypo
th
e
s
i
s
:f
em
a
l
e
seng
ag
einEPC
sw
i
thm
a
l
e
so
fh
igh
e
rqu
a
l
i
tyth
anth
e
i
rso
c
i
a
lm
a
t
e
sto

17

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e
s
i
s (2017
)

ob
t
a
inf
avou
r
ab
l
eg
en
e
sfo
rth
e
i
ro
f
f
sp
r
ing
.F
em
a
l
e
sm
aya
s
s
e
s
sm
al
equ
a
l
i
tyu
s
ing
ph
eno
typ
i
cs
ign
a
l
s(
e
.g
.
,p
lum
ag
eo
rn
am
en
t
a
t
ion
,song
,o
rb
eh
av
iou
r
)th
a
tconv
ey
in
fo
rm
a
t
ionabou
tphy
s
i
c
a
lcond
i
t
ion(
e
.g
.
,nu
t
r
i
t
ion
a
ls
t
a
t
e
)and
/o
rg
en
e
t
i
cqu
a
l
i
ty
.Fo
r
ex
amp
l
e
,f
em
a
l
eb
lu
et
i
t
s(
C
yan
i
s
t
e
sca
e
ru
l
eu
s)eng
ag
einEPC
sw
i
tho
ld
e
r
,l
a
rg
e
rm
a
l
e
s
(K
emp
en
a
e
r
se
ta
l
.1997
)andm
a
l
e
sw
i
thb
r
igh
t
e
ru
l
t
r
av
io
l
e
tb
lu
ep
lum
ag
e(K
emp
en
a
e
r
se
t
a
l
.1992
)
,wh
i
l
ef
em
a
l
eb
l
a
ck
c
app
edch
i
ck
ad
e
e
seng
ag
einEPC
sw
i
thmo
r
edom
in
an
tm
a
l
e
s
(Sm
i
th1988
;O
t
t
e
re
ta
l
.1994
,1998
;M
enn
i
l
le
ta
l
.2004
)
.
H
ab
i
t
a
tu
rb
an
i
z
a
t
ionc
ana
f
f
e
c
tfoodav
a
i
l
ab
i
l
i
tyandin
t
e
r
sp
e
c
i
f
i
cin
t
e
r
a
c
t
ion
s
,and
con
s
equ
en
t
ly
,m
ayimp
a
c
taf
em
a
l
e
’
sl
ik
e
l
ihoodtobo
ths
e
ekanda
c
c
ep
tEPC
s
.Ex
t
r
ap
a
i
r
p
a
t
e
rn
i
tyh
a
sb
e
enr
e
l
a
t
edtoh
ab
i
t
a
tqu
a
l
i
tyino
th
e
rsongb
i
rdsp
e
c
i
e
s
.Inhou
s
esp
a
r
row
s
,
exp
e
r
im
e
n
t
a
l
lyin
c
r
e
a
s
edfoodav
a
i
l
ab
i
l
i
tyr
e
su
l
t
edinp
a
i
r
ssp
end
ingmo
r
et
im
etog
e
th
e
ra
t
th
en
e
s
t
,wh
i
ch
,intu
rn
,l
e
dtoaf
iv
efo
ldr
edu
c
t
ioninEPPr
a
t
e
s(V
á
c
l
ave
ta
l
.2003
)
.In
sup
e
rbs
t
a
r
l
ing
s(
Lamp
ro
to
rn
i
ssup
e
rbu
s)
,Rub
en
s
t
e
in(2007
)foundEPPtob
el
e
s
sp
r
ev
a
l
en
t
inh
igh
e
r
qu
a
l
i
tyt
e
r
r
i
to
r
i
e
s(
i
.
e
.
,g
r
e
a
t
e
rv
eg
e
t
a
t
ioncov
e
randp
r
eyav
a
i
l
ab
i
l
i
ty
)
.A
sth
eau
tho
r
sugg
e
s
t
s
,g
r
e
a
t
e
rp
r
eyav
a
i
l
ab
i
l
i
tyi
nh
igh
qu
a
l
i
tyt
e
r
r
i
to
r
i
e
sm
ayl
im
i
tth
ed
i
s
t
an
c
ef
em
a
l
e
s
n
e
edtot
r
av
e
ltofo
r
ag
e
,andcon
s
equ
en
t
ly
,d
e
c
r
e
a
s
eh
e
rp
rob
ab
i
l
i
tyo
fen
coun
t
e
r
inganEP
m
a
l
e
.Incon
t
r
a
s
t
,s
e
r
in(
S
e
r
inu
ss
e
r
inu
s)n
e
s
t
sint
e
r
r
i
to
r
i
e
sw
i
thg
r
e
a
t
e
rfoodav
a
i
l
ab
i
l
i
tya
r
e
mo
r
el
ik
e
lytocon
t
a
inEPOth
ann
e
s
t
sinpoo
rqu
a
l
i
tyh
ab
i
t
a
t(Ho
i
L
e
i
tn
e
re
ta
l
.1999
).Th
e
au
tho
r
spo
s
tu
l
a
t
ef
em
a
l
e
sonh
igh
qu
a
l
i
tyt
e
r
r
i
to
r
i
e
sm
ayb
einb
e
t
t
e
rcond
i
t
ionandmo
r
eab
l
e
tor
e
s
i
s
tm
a
l
em
a
t
egu
a
rd
inge
f
fo
r
t
s
,andthu
s
,m
ayh
av
eg
r
e
a
t
e
roppo
r
tun
i
tyeng
ag
einEPC
s
(Ho
i
L
e
i
tn
e
re
ta
l
.1999
).
Fo
rspo
t
t
edtowh
e
e
s(
P
ip
i
loma
cu
la
tu
s)b
r
e
ed
inginu
rb
anp
a
rk
s
,p
a
rkedg
e
sa
r
es
i
t
e
s
o
fh
ighfoodabund
an
c
edu
etoan
th
ropog
en
i
cfoodsou
r
c
e
s(
e
.g
.
,b
i
rdf
e
ed
e
r
s
)
,wh
i
l
ein
t
e
r
io
r
h
ab
i
t
a
ti
scomp
a
r
a
t
iv
e
lylow
e
r
qu
a
l
i
ty(Sm
i
the
ta
l
.2
016
).I
fEPCr
a
t
e
sd
e
c
l
in
ew
i
th
18

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

in
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anth
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i
the
ta
l
.(2016
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19

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

2
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20

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
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s (2017
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21

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
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2
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22

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
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23

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

w
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24

E
r
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aS
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rud

M
.S
c
.Th
e
s
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s (2017
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ly
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ing2014a
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ing
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in
a
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er
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to
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r
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p
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r
t
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fEPOinh
i
sn
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s
t
,w
econ
s
t
ru
c
teds
im
i
l
a
rmod
e
l
s
,bu
tu
s
edth
enumb
e
ro
fEPOin
th
en
e
s
ta
sth
er
e
spon
s
ev
a
r
i
ab
l
eandto
t
a
lnumb
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ro
fo
f
f
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r
inginth
en
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s
ta
sth
eb
inom
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in
a
to
r
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skhowf
em
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l
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f
lu
en
c
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sh
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rp
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andome
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lu
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c
edth
ep
r
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s
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c
eandp
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r
t
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fEPOinn
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s
t
s
,w
e
con
s
t
ru
c
t
eds
im
i
l
a
rmod
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l
su
s
ingou
rh
ab
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a
tandv
eg
e
t
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t
ionind
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sa
sth
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r
ed
i
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to
rv
a
r
i
ab
l
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s
,
andf
em
a
l
eid
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t
i
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sar
andome
f
f
e
c
t
.F
in
a
l
ly
,toa
skwh
e
th
e
rp
a
r
en
t
a
lp
rov
i
s
ion
ingw
a
s
25

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

ad
ju
s
t
edinr
e
spon
s
etoth
ep
r
e
s
en
c
eo
rp
ropo
r
t
iono
fEPOinan
e
s
t
,w
econ
s
t
ru
c
t
ed
g
en
e
r
a
l
i
z
edl
in
e
a
rmod
e
l
su
s
ingp
a
r
en
t
a
lp
rov
i
s
ion
ingr
a
t
e(
t
r
ip
s
/h
r
/
ch
i
ck
)a
sth
er
e
spon
s
e
v
a
r
i
ab
l
e
,andn
e
s
tp
a
t
e
rn
i
tyandth
ep
ropo
r
t
iono
fEPOinth
en
e
s
ta
sth
ep
r
ed
i
c
to
rv
a
r
i
ab
l
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s
.
W
eex
c
lud
edth
es
ing
l
ec
a
s
eo
fr
en
e
s
t
inginth
e2016b
r
e
ed
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e
a
son
.W
eth
eni
so
l
a
tedon
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s
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s
t
scon
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a
in
ingEPO
,andr
e
r
anth
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c
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ingon
lyth
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r
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fEPOinth
e
n
e
s
ta
sap
r
ed
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to
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ab
l
e
.

2
.4
. R
e
su
l
t
s
2
.4
.1
.G
en
e
ra
lpa
t
t
e
rn
so
fpa
r
en
tag
e
Ov
e
r
a
l
l
,17
.8%(46
/258
)o
fo
f
f
sp
r
ingw
e
r
es
i
r
edbyanEPm
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l
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,and44
.4%(20
/45
)ofn
e
s
t
s
con
t
a
in
edEPO(F
igu
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e2
.1
)
.Th
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e
r
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t
ag
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fEPOw
i
th
inn
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s
t
sw
i
thm
ix
edp
a
r
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ag
e
r
ang
edf
rom12
.5%(1
/8o
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f
sp
r
ingEP
)to100%(4
/4o
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f
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r
ingEP
)
.W
ew
e
r
eab
l
eto
con
f
id
en
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lyid
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t
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fys
ev
enEPm
a
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sf
roms
ix(30
.0%
)o
fth
em
ix
ed
p
a
t
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rn
i
tyn
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s
,w
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thtwo
EPm
a
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ss
i
r
ingEPOw
i
th
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roodinon
ec
a
s
e
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thth
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i
a
lm
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l
eandEPm
a
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e
r
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rfou
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e
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/EPm
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s
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ecomp
a
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r
e
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e
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s
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r
ew
a
sag
r
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2
=0
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;2014v
s
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=1
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;2015v
s
.
in2016(2014v
s
.2015
:χ
2
2016
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=2
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)
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l
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f
f
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ingw
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efoundtob
eth
eg
en
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co
f
f
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r
ingo
fth
e
i
r

pu
t
a
t
iv
emo
th
e
r
.

26

E
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i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

2
.4
.2
. Cond
i
t
ionande
x
t
rapa
i
rpa
t
e
rn
i
t
y
W
ed
idno
tf
inde
i
th
e
rf
em
a
l
eag
eo
rbodycond
i
t
iontoin
f
lu
en
c
eaf
em
al
e
’
sl
ik
e
l
ihoodto
eng
ag
einEPC
so
rth
ep
ropo
r
t
iono
fEPOinh
e
rn
e
s
t(
a
l
lp>0
.21;T
ab
l
e2
.2
)
. Wh
enw
e
con
s
id
e
r
edm
a
l
econd
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t
ion
,w
ed
idno
tf
indag
e
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t
ion
,o
rcho
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sch
a
r
a
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t
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t
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in
f
lu
en
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’
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ik
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ihoodo
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s
ingp
a
t
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rn
i
ty(
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l
lp>0
.44;T
ab
l
e2
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)
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ea
l
sod
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a
r
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r
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e
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ab
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)
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id
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e
r
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t(
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ab
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2
.4
.3
. So
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ia
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l
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s
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t
ra
pa
i
rma
l
ecompa
r
i
son
s
Wh
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edth
econd
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t
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tytoth
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.24)o
rag
e(
W =1
.73
,p=0
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)b
e
tw
e
enth
e
d
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f
f
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r
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c
einbodycond
i
t
i
on(t
4=1
two
,bu
tth
esm
a
l
ls
amp
l
es
i
z
e(
n=5
)fo
rth
e
s
ecomp
a
r
i
son
sr
e
su
l
t
edinlow
e
rpow
e
r
.Ina
l
l
f
iv
eso
c
i
a
lm
a
l
e
/EPm
a
l
ep
a
i
r
s
,th
eEPm
a
l
ew
a
so
fequ
iv
a
l
en
tag
etoth
eso
c
i
a
lm
a
l
e(ASY
),o
ro
ld
e
r
th
an(ASYv
s
.SY
,n=3
).Inth
etwocomp
a
r
i
son
swh
e
r
eth
em
a
l
e
s
v
s
.AS
Y
,n=2
w
e
r
eo
fequ
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l
en
tag
e
,twom
a
l
e
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ld
ingn
e
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r
ingt
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to
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i
e
sonth
eThomp
sonR
iv
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r
s
Un
iv
e
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s
i
tyc
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ss
i
r
edEPOine
a
cho
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e
r
’
sn
e
s
t
s(
i
.
e
.
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thg
a
in
edp
a
t
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ty
,bu
ta
l
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th
lo
s
tp
a
t
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rn
i
tytoon
eano
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e
r
)
.

2
.4
.4
. Hab
i
ta
te
f
f
e
c
t
s
W
efoundn
e
i
th
e
rth
eh
ab
i
t
a
tind
exno
rv
eg
e
t
a
t
ioncompo
s
i
t
iono
fn
e
s
t
ingh
ab
i
t
a
ttoh
av
ean
e
f
f
e
c
tone
i
th
e
rth
ep
r
e
s
en
c
eo
rp
ropo
r
t
iono
fEPOinan
e
s
t(
a
l
lp>0
.09
;T
ab
l
e2
.2
,F
igu
r
e
2
.2)
.
27

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

2
.4
.5
.E
x
t
rapa
i
rpa
t
e
rn
i
t
yandpa
r
en
ta
lca
r
e
Wh
i
l
ene
s
t
scon
t
a
in
ingEPOt
end
ed
,ov
e
r
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l
,tob
ep
rov
i
s
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eda
talow
e
rr
a
t
eth
antho
s
e
w
i
thou
t
,(
thoughno
ts
ign
i
f
i
c
an
t
ly
:p=0
.06
;T
ab
l
e2
.3
)
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i
s
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ingr
a
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r
epo
s
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ly
r
e
l
a
t
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r
t
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fo
f
f
s
p
r
inginan
e
s
tth
a
tw
e
r
eEP
O(
p=0
.007
;T
ab
l
e2
.3
,F
igu
r
e
2
.3)
. Wh
enw
ei
so
l
a
t
edn
e
s
t
scon
t
a
in
ingEPO
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er
e
l
a
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sh
ipb
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tw
e
enp
rov
i
s
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ingr
a
t
eand
p=0
.006
;T
ab
l
e2
.3
,F
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r
e2
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)
.
th
ep
ropo
r
t
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fEPOinth
en
e
s
tr
em
a
in
eds
ign
i
f
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c
an
t(

2
.5
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s
cu
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s
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Ex
t
r
ap
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s
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a
s
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t
i
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e
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s
t
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a
t
edinmoun
t
a
inch
i
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e
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s
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e
r
e
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r
e
s
en
tev
id
en
c
eth
a
tmoun
t
a
inch
i
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e
e
s
a
l
sof
r
equ
en
t
lyeng
ag
einEPC
s
:EPOw
e
r
efoundina
lmo
s
t50%o
fn
e
s
t
sandr
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p
r
e
s
en
t
ed
a
lmo
s
t20%o
fo
f
f
sp
r
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.Th
e
s
ew
e
r
esom
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a
tsu
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r
i
s
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lyh
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r
equ
en
c
i
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s
,a
sr
a
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so
f
EPPinth
ec
lo
s
e
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e
l
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t
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l
a
ckc
app
edch
i
ck
ad
e
et
enda
round30%o
fn
e
s
t
sand10
15%o
f
o
f
f
sp
r
ing(O
t
t
e
re
ta
l
.1998
;R
am
s
aye
ta
l
.2000
;M
enn
i
l
le
ta
l
.2004
)
.How
ev
e
r
,s
im
i
l
a
r
ly
h
igh
,andev
enh
igh
e
r
,r
a
t
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so
fEPPtowh
a
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eob
s
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r
v
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l
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fn
e
s
t
s
;D
e
lh
eye
ta
l
.2007
)andg
r
e
a
tt
i
t(
Pa
ru
sma
jo
r,40% o
fn
e
s
t
s
;Lub
juhn
e
ta
l
.1999
)
.Inadd
i
t
ion
,h
ighr
a
t
e
so
fEPPh
av
eb
e
enob
s
e
rv
edinb
l
a
ck
c
app
edch
i
ck
ad
e
e
s
b
r
e
ed
inginhyb
r
idzon
e
sw
i
thC
a
ro
l
in
ach
i
ck
ad
e
e
s(
Po
e
c
i
l
eca
ro
l
in
en
s
i
s)(55
.6%o
fn
e
s
t
s
;
R
eud
inke
ta
l
.2006
)andmoun
t
a
inch
i
ck
ad
e
e
s(62
.5%o
fn
e
s
t
sG
r
av
ae
ta
l
.2012
)
.
Th
ep
r
e
s
en
c
eo
fEPPinP
a
r
id
a
esp
e
c
i
e
sh
a
so
f
t
enb
e
ena
t
t
r
ibu
t
edtothegoodg
en
e
s
hypo
th
e
s
i
s
:f
em
a
l
e
seng
ag
einEPC
stoob
t
a
inmo
r
ef
avou
r
ab
l
eg
en
e
sfo
rth
e
i
ro
f
f
sp
r
ingth
an
th
e
i
rso
c
i
a
lm
a
t
ec
anp
rov
id
etoin
c
r
e
a
s
eth
e
i
rownf
i
tn
e
s
s
.Inb
l
a
ck
c
app
edch
i
ck
ad
e
e
s
,
f
em
a
l
e
seng
ag
einEPC
sw
i
thm
a
l
e
so
fh
igh
e
rdom
in
an
c
er
ankth
anth
e
i
rso
c
i
a
lm
a
l
e(O
t
t
e
re
t
28

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

a
l
.1998
;M
enn
i
l
le
ta
l
.2004
)
.Inb
lu
et
i
t
s
,s
ev
e
r
a
lm
e
a
su
r
e
so
fcond
i
t
ionapp
e
a
rtoin
f
lu
en
c
e
th
ed
e
c
i
s
ion
so
ff
em
a
l
e
s
,w
i
thl
a
rg
e
rm
a
l
e
s(K
emp
en
a
e
r
se
ta
l
.1997
)
,o
ld
e
rm
a
l
e
s
(K
emp
en
a
e
r
se
ta
l
.1997
)
,m
a
l
e
sw
i
thg
r
e
a
t
e
rsongou
tp
u
t(K
emp
en
a
e
r
se
ta
l
.1997
)
,andm
a
l
e
whob
eg
ins
ing
inge
a
r
l
i
e
r(Po
e
s
e
le
ta
l
.2006
)b
e
ingsough
ta
sEPs
i
r
e
s
.H
e
r
e
,w
ea
sk
edhow
m
a
l
econd
i
t
ion
,a
sm
e
a
su
r
edbyag
e
,w
e
igh
tr
e
l
a
t
iv
etobodys
i
z
e
,andd
awncho
ru
sou
tpu
t
in
f
lu
en
c
eh
i
sl
ik
e
l
ihoodtolo
s
ep
a
t
e
rn
i
ty
.W
ed
idno
tf
indanym
e
a
su
r
e
so
fm
a
l
econd
i
t
ionto
p
r
ed
i
c
tth
ep
r
e
s
en
c
eo
fEPOinh
i
sn
e
s
t
,no
rd
idw
ef
indanys
ign
i
f
i
c
an
tr
e
l
a
t
ion
sh
ipb
e
tw
e
en
m
a
l
econd
i
t
ionandth
ep
ropo
r
t
iono
fEPOinh
i
sn
e
s
t
.W
ed
id
,how
ev
e
r
,f
indan
e
a
r
s
ign
i
f
i
c
an
t(
p=0
.07)e
f
f
e
c
to
fm
a
l
eag
eandth
ep
ropo
r
t
iono
fEPOinh
i
sn
e
s
t
.O
ld
e
r
,AS
Y
m
a
l
e
st
end
edtor
a
i
s
ef
ew
e
rEPO
,af
ind
ingcon
s
i
s
t
en
tw
i
thth
egoodg
en
e
shypo
th
e
s
i
s
.In
add
i
t
ion
,w
efoundth
a
tina
l
lc
a
s
e
swh
e
r
ebo
thth
eso
c
i
a
lm
a
l
eandEPm
a
l
ew
e
r
eknown
,th
e
EPm
a
l
ew
a
se
i
th
e
ro
fequ
iv
a
l
en
tag
etoth
eso
c
i
a
lm
a
l
eo
ro
ld
e
r
.Inth
ec
a
s
e
swh
e
r
eth
em
a
l
e
s
w
e
r
eo
fequ
iv
a
l
en
tag
e
,bo
thm
a
l
e
sw
e
r
eASY
,thoughth
e
i
rag
einy
e
a
r
sw
a
sno
tknown
.
Tog
e
th
e
r
,th
e
s
ean
e
cdo
t
e
ssugg
e
s
tf
em
a
l
emoun
t
a
inch
i
ck
ad
e
e
seng
ag
einEPC
sw
i
thadu
l
t
m
a
l
e
s(5
/5c
a
s
e
s
)
,andm
a
l
e
so
ld
e
rth
anth
e
i
rso
c
i
a
lm
a
l
e(3
/5con
f
i
rm
edc
a
s
e
s
)–f
ind
ing
s
ag
a
incon
s
i
s
t
en
tw
i
thth
egoodg
en
e
shypo
th
e
s
i
s
.
W
efoundmoun
t
a
inch
i
ck
ad
e
e
sad
ju
s
tp
rov
i
s
ion
inge
f
fo
r
tinr
e
l
a
t
ionton
e
s
tp
a
t
e
rn
i
ty
.
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t
rongpo
s
i
t
iv
er
e
l
a
t
ion
sh
ipb
e
tw
e
enp
rov
i
s
ion
inge
f
fo
r
tandth
ep
ropo
r
t
iono
f
EPOinan
e
s
t.B
e
c
au
s
ei
tw
a
sd
i
f
f
i
cu
l
ttod
i
s
c
e
rnind
iv
idu
a
lid
en
t
i
tyinou
rn
e
s
tw
a
t
chv
id
eo
s
,
w
ea
s
s
e
s
s
edp
rov
i
s
ion
ingr
a
t
ea
sth
eco
l
l
e
c
t
iv
ep
rov
i
s
ion
inge
f
fo
r
to
fbo
thth
em
a
l
eand
f
em
a
l
e
.I
ff
em
a
l
e
sa
r
ead
ju
s
t
ingth
e
i
rp
rov
i
s
ion
ing
,th
i
scou
ldp
r
e
s
en
tac
a
s
eo
fd
i
f
f
e
r
en
t
i
a
l
a
l
lo
c
a
t
ion(Bu
r
l
ey1986
)
,w
i
thf
em
a
l
e
sin
c
r
e
a
s
ingp
rov
i
s
ion
ingb
e
c
au
s
eth
ey‘know
’th
e
r
ea
r
e
som
epo
t
en
t
i
a
l
lyh
igh
e
r
qu
a
l
i
tyEPOinth
e
i
rn
e
s
t
.A
l
t
e
rn
a
t
iv
e
ly
,i
fm
a
l
e
sa
r
eth
eon
e
s
in
c
r
e
a
s
ingth
e
i
rp
a
r
en
t
a
le
f
fo
r
t
,th
i
scou
ldb
eas
i
tu
a
t
ionl
ik
einhou
s
ef
in
ch
e
swh
e
r
e
,insom
e
29

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

popu
l
a
t
ion
s
,y
e
l
l
owm
a
l
e
s(
l
e
s
sp
r
e
f
e
r
r
edbyf
em
a
l
e
s
)p
rov
i
s
iono
f
f
sp
r
inga
tah
igh
e
rr
a
t
e
(B
ady
a
evandH
i
l
l2002
)
.How
ev
e
r
,no
tknow
ingwhoi
sdo
ingth
ep
rov
i
s
ion
ingm
ak
e
si
t
d
i
f
f
i
cu
l
ttoin
t
e
rp
r
e
tth
ep
a
t
t
e
rn
.
I
ncon
t
r
a
s
ttomu
l
t
ip
l
eo
th
e
rs
tud
i
e
s
,w
ef
a
i
l
edtof
indas
ign
i
f
i
c
an
te
f
f
e
c
to
fh
ab
i
t
a
ton
EPPinmoun
t
a
inch
i
ck
ad
e
e
s
.S
ev
e
r
a
ls
tud
i
e
sinv
e
s
t
ig
a
t
ingh
ab
i
t
a
tandEPPh
av
econ
s
id
e
r
ed
d
i
f
f
e
r
en
c
e
sinfoodav
a
i
l
ab
i
l
i
tyino
th
e
rw
i
s
es
im
i
l
a
rh
ab
i
t
a
ta
sth
ed
e
t
e
rm
in
an
to
fh
ab
i
t
a
t
qu
a
l
i
ty(Ho
i
L
e
i
tn
e
re
ta
l
.1999
;V
á
c
l
ave
ta
l
.2003
;Rub
en
s
t
e
in2007
)
.H
e
r
e
,w
ea
sk
edhow
EPPv
a
r
i
eda
longag
r
ad
i
en
tf
romru
r
a
ltou
rb
anh
ab
i
t
a
t
.Un
l
ik
eo
th
e
rs
tud
i
e
s
,w
eh
adno
d
i
r
e
c
tm
e
a
su
r
eo
fh
ab
i
t
a
tqu
a
l
i
ty(
e
.g
.
,foodav
a
i
l
ab
i
l
i
ty
)
.How
ev
e
r
,u
rb
anh
ab
i
t
a
tapp
e
a
r
sto
b
eb
e
t
t
e
rqu
a
l
i
tyth
anru
r
a
lh
ab
i
t
a
ttoth
emoun
t
a
inch
i
ck
ad
e
e
sinou
rs
tudypopu
l
a
t
ion
(M
a
r
in
ie
ta
l
.2017
)
,p
r
e
sum
ab
lyb
e
c
au
s
eo
fg
r
e
a
t
e
rfoodav
a
i
l
ab
i
l
i
tydu
etoth
ep
r
e
s
en
c
eo
f
b
i
rdf
e
ed
e
r
sandg
r
e
a
t
e
rd
e
c
iduou
st
r
e
econ
t
en
t
.S
t
i
l
l
,w
ed
idn
o
tf
indEPPtob
er
e
l
a
t
edtoth
e
h
ab
i
t
a
tind
exo
rv
eg
e
t
a
t
ioncompo
s
i
t
iono
fn
e
s
t
ings
i
t
e
s
,ind
i
c
e
sth
a
tw
e
r
ed
e
r
iv
edtod
e
s
c
r
ib
e
h
ab
i
t
a
tu
rb
an
i
z
a
t
ionandqu
a
l
i
ty
,andv
eg
e
t
a
t
ioncompo
s
i
t
ionandfoodav
a
i
l
ab
i
l
i
ty
,
r
e
sp
e
c
t
iv
e
ly
.
O
th
e
rs
tud
i
e
sw
i
ths
i
t
e
sbord
e
r
ingu
rb
anandsubu
rb
ann
e
ighbou
rhood
sh
av
efound
e
f
f
e
c
t
so
fu
rb
an
i
z
a
t
iononEPP
.Fo
rspo
t
t
edtowh
e
e
sb
r
e
ed
inginu
rb
anp
a
rk
s
,EPPr
a
t
e
sa
r
e
h
igh
e
s
tinin
t
e
r
io
rh
ab
i
t
a
tandu
rb
an
bo
rd
e
r
ingedg
eh
ab
i
t
a
t
,andlow
e
s
ta
tin
t
e
rm
ed
i
a
t
e
d
i
s
t
an
c
e
sf
romth
eedg
e(Sm
i
the
ta
l
.2016
)
.Inth
e
s
ep
a
rk
s
,edg
eh
ab
i
t
a
ti
scomp
a
r
a
t
iv
e
ly
b
e
t
t
e
rqu
a
l
i
tyth
anin
t
e
r
io
rh
ab
i
t
a
t
,a
sth
e
r
ei
sg
r
e
a
t
e
rfoodabund
an
c
ea
tedg
e
sdu
eto
an
th
ropog
en
i
cfoodsou
r
c
e
s(Sm
i
the
ta
l
.2016
)
.A
ssu
ch
,th
eh
ighp
r
ev
a
l
en
c
eo
fEPPin
in
t
e
r
io
rt
e
r
r
i
to
r
i
e
sw
a
sexp
e
c
t
ed
,bu
tth
es
am
eob
s
e
rv
a
t
ioninedg
et
e
r
r
i
to
r
i
e
sw
a
sno
t
.Th
e
au
tho
r
ssugg
e
s
tadd
i
t
ion
a
lfoodsou
r
c
e
sinedg
eh
ab
i
t
a
td
r
ewind
iv
idu
a
l
sf
romth
e
su
r
round
inglow
e
r
qu
al
i
tyh
ab
i
t
a
t
,r
e
su
l
t
inginh
igh
e
rd
en
s
i
t
i
e
so
ft
r
an
s
i
en
tb
i
rd
sinth
eedg
e
30

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

h
ab
i
t
a
tandin
c
r
e
a
s
edin
s
t
an
c
e
so
fEPC
s(Sm
i
the
ta
l
.2016
)
.Inb
lu
et
i
t
s
,K
emp
en
a
e
r
se
ta
l
.
(2010
)founda
r
t
i
f
i
c
i
a
ln
igh
tl
igh
t
ing(
i
.
e
.
,s
t
r
e
e
tl
igh
t
s
)insubu
rb
anh
ab
i
t
a
tin
f
lu
en
c
e
sEPP
.
Comp
a
r
edtom
a
l
e
sw
i
tht
e
r
r
i
to
r
i
e
sinin
t
e
r
io
rfo
r
e
s
t
,m
a
l
e
sinedg
eh
ab
i
t
a
tbo
rd
e
r
ingl
igh
t
ed
subu
rb
ann
e
ighbou
rhood
sw
e
r
eno
tinanyb
e
t
t
e
rcond
i
t
ion
,bu
tw
e
r
emo
r
esu
c
c
e
s
s
fu
la
t
g
a
in
ingp
a
t
e
rn
i
tyino
th
e
rn
e
s
t
s(K
emp
en
a
e
r
se
ta
l
.2010
)
.M
a
l
e
sinl
igh
t
edt
e
r
r
i
to
r
i
e
sa
l
so
b
eg
ans
ing
ingea
r
l
i
e
r(K
emp
en
a
e
r
se
ta
l
.2010
)
.Inn
a
tu
r
a
l
,fo
r
e
s
th
ab
i
t
a
t
,f
em
a
l
eb
lu
et
i
t
s
eng
ag
einEPC
sw
i
the
a
r
lys
ing
ingm
a
l
e
s
,sugg
e
s
t
ingth
et
im
ingo
fd
awns
ing
ingm
ayb
ean
ind
i
c
a
to
ro
fm
a
l
equ
a
l
i
ty(K
emp
en
a
e
r
se
ta
l
.1992
)
.Thu
s
,f
em
a
l
e
sm
ayh
av
ep
e
r
c
e
iv
ede
a
r
ly
s
ing
ingm
a
l
e
sinsubu
rb
anh
ab
i
t
a
ta
sb
e
ingh
ighqu
a
l
i
tyand
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s
equ
en
t
ly
,sough
tth
e
s
e
m
a
l
e
sa
sEPp
a
r
tn
e
r
s
.
L
ik
einou
rs
tudypopu
l
a
t
ion(M
a
r
in
ie
ta
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.2017
)
,K
emp
en
a
e
rse
ta
l
.(2010
)found
f
em
a
l
e
sn
e
s
t
inginsubu
rb
ant
e
r
r
i
to
r
i
e
sb
eg
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ay
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se
a
r
l
i
e
r
.F
em
a
l
e
sshou
ldt
im
e
b
r
e
ed
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a
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e
s
t
l
ingfoodd
em
anda
l
ign
sw
i
thp
e
akfoodav
a
i
l
ab
i
l
i
ty
,how
ev
e
r
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s
th
eau
tho
r
ssugg
e
s
t
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a
r
l
i
e
rl
ay
ingm
ayh
av
el
edtoam
i
sm
a
t
chb
e
tw
e
enth
etwoinsubu
rb
an
h
ab
i
t
a
t(K
emp
en
a
e
r
se
ta
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.2010
)
.Thu
s
,r
a
th
e
rth
anind
i
c
a
t
ingb
e
t
t
e
rh
ab
i
t
a
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a
l
i
ty
,a
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e
h
adsp
e
cu
l
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t
ed
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r
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lu
t
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t
i
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t
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rb
anh
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i
t
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tm
ayb
eam
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l
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ap
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eb
eh
av
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r
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ig
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t
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tu
r
e
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rb
anh
ab
i
t
a
t(
e
.g
.
,a
r
t
i
f
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c
i
a
ll
igh
t
ing
)
.Inv
e
s
t
ig
a
t
ion
so
fth
er
e
l
a
t
iv
e
t
im
ingo
fb
r
e
ed
inginr
e
l
a
t
iontoin
s
e
c
tabund
an
c
ep
e
ak
sine
i
th
e
rh
ab
i
t
a
twou
ldh
av
etob
e
condu
c
t
edtod
i
s
c
e
rnwh
e
th
e
rth
e
s
em
i
sm
a
t
ch
e
so
c
cu
r
.Tog
e
th
e
r
,th
e
s
eex
amp
l
e
si
l
lu
s
t
r
a
t
eth
e
comp
l
exdyn
am
i
c
so
fu
rb
anh
ab
i
t
a
t
,andsugg
e
s
tth
ed
i
f
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e
r
en
c
e
sw
eh
av
ep
r
ev
iou
s
lyob
s
e
rv
ed
b
e
tw
e
enmoun
t
a
inch
i
ck
ad
e
e
sinu
rb
anandru
r
a
lh
ab
i
t
a
t(
s
e
eM
a
r
in
ie
ta
l
.2017
)m
ay
r
ep
r
e
s
en
tmo
r
eth
ans
imp
lyd
i
f
f
e
r
en
c
e
sinh
ab
i
t
a
tqu
a
l
i
ty
.

2
.6
. L
i
t
e
ra
tu
r
ec
i
t
ed
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t
r
a
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egy
:d
i
s
t
in
c
t
r
ep
rodu
c
t
iv
et
a
c
t
i
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sa
s
so
c
i
a
t
edw
i
the
l
abo
r
a
t
iono
fp
lum
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a
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sh
ip
sandhyb
r
id
i
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a
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ionb
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tw
e
enb
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a
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c
app
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andmoun
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a
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:at
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rin
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.A
.(2016
)
.Sp
a
t
i
alp
a
t
t
e
rn
so
fex
t
r
a
ip
i
loma
cu
la
tu
sinu
rb
anp
a
rk
s
.— J
.Av
i
anB
io
l
.47
:
p
a
i
rp
a
t
e
rn
i
tyfo
rspo
t
t
edtowh
e
e
sP
001
009
Sm
i
th
,S
.M
.(1988
)
.Ex
t
r
ap
a
i
rcopu
l
a
t
ion
sinb
l
a
ckc
app
edch
i
ck
ad
e
e
s
:th
ero
l
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fth
e
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em
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l
e
.— B
eh
av
iou
r107
:1523
.
V
á
c
l
av
,R
.
,Ho
i
,H
.&B
lomqv
i
s
t
,D
.(2003
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.Foodsupp
l
em
en
t
a
t
iona
f
f
e
c
t
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t
r
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a
i
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r
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cu
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eh
av
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l
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ao
,C
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.andL
i
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.H
.(2005
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l
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lu
s)
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— Mo
l
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co
l
.No
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s5
:439442
.

33

E
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M
.S
c
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2
.7
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34

E
r
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M
.S
c
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T
ab
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35

E
r
i
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aS
.Bond
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rud

M
.S
c
.Th
e
s
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s (2017
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T
ab
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36

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

2
.8
. F
igu
r
e
s

Pe
r
cen
tageo
fne
s
t
s

100%
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15

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13

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37

-4 -3 -2 -1

U
rban

0

1

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ra
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0
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l
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ty
.

38

M
.S
c
.Th
e
s
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s(2017
)

2
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)
.

39

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

CHAPTER3
:EFFECTSOFPARENTALCOND
IT
IONANDURBAN
IZAT
IONON
SEXALLOCAT
ION
3
.1
. Ab
s
t
ra
c
t
Wh
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er
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lu
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s
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t
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sd
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f
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ldb
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t
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.

3
.2
. In
t
rodu
c
t
ion
S
exa
l
lo
c
a
t
ionth
eo
ryp
r
ed
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c
t
sf
em
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l
e
sshou
ldb
i
a
sth
es
exr
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t
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fth
e
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ro
f
f
sp
r
inginr
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spon
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tod
i
f
f
e
r
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e
sinth
er
ep
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c
t
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ev
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lu
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fson
sandd
augh
t
e
r
s(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;
Ch
a
rnov1982
).O
f
f
sp
r
ingr
ep
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c
t
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ev
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lu
em
ayv
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rya
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e
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l
to
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t
r
in
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i
cf
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c
to
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s
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.
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a
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ab
i
t
a
t
qu
a
l
i
ty
)(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;Bu
r
l
ey1981
)
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e
r
e
fo
r
e
,f
em
a
l
e
sshou
ldb
i
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roods
ex
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spon
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a
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n
t
a
l

40

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

cond
i
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ionandh
ab
i
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a
tqu
a
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i
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s(
r
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av
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i
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tu
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sgov
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an
c
eh
i
e
r
a
r
ch
i
e
s(M
cC
a
l
l
ume
ta
l
.1999
;R
a
t
c
l
i
f
f
ee
ta
l
.2007
)
.Inb
l
a
ck
c
app
ed
ch
i
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e
s
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r
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a
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c
l
i
f
f
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ta
l
.2007
)
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i
a
lm
a
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e
s(O
t
t
e
randR
a
t
c
l
i
f
f
e1996
;R
am
s
aye
ta
l
.2000
)
a
ndEP
p
a
r
tn
e
r
s(O
t
t
e
re
ta
l
.1994
,1998
)
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a
l
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ss
ign
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lth
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i
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t
a
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sth
roughcond
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a
l
i
ty(O
t
t
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re
ta
l
.1997
;M
enn
i
l
le
t
a
l
.2003
)
.Thu
s
,f
em
a
l
e
sm
ayb
eab
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eg
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t
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se
ta
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.2005
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lu
et
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l
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(2014
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ayh
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t
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es
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a
l
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rp
a
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am
s
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ta
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.2003
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i
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ly
41

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

co
r
r
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l
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edw
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thh
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/qu
a
l
i
ty(
e
.g
.
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l
l
ik
e
r
e
ta
l
.1999
;Sh
e
ldone
ta
l
.1999
;S
t
au
s
se
ta
l
.2005
)ino
th
e
rP
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i
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thCh
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s(1982
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fs
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42

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M
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.Th
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1998
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43

E
r
i
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aS
.Bond
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rud

M
.S
c
.Th
e
s
i
s (2017
)

b
r
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rud

M
.S
c
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s
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s (2017
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45

E
r
i
c
aS
.Bond
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rud

M
.S
c
.Th
e
s
i
s (2017
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46

E
r
i
c
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rud

M
.S
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.Th
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s
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2
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x
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lytob
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sth
eh
ab
i
t
a
tb
e
c
am
e
47

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

mo
r
eu
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an
i
z
ed
.W
ed
idno
tf
indf
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l
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fs
exa
l
lo
c
a
t
ion(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;Ch
a
rnov1982
)
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e
p
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ed
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c
t
edu
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ab
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e
i
rru
r
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rp
a
r
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s(M
a
r
in
ie
ta
l
.2017
)
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s
t
ingu
rb
anh
ab
i
t
a
tm
ay
p
rov
id
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s
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sp
ropo
s
edby M
a
r
in
ie
ta
l
.(2017
)
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rb
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e
s
ts
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t
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ay
p
rov
id
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r
e
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l
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t
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inch
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roughth
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r
e
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i
rdf
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s
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rb
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ab
i
t
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ti
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s
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r
e
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rd
e
c
iduou
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r
e
eabund
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e
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l
though
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t
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inch
i
ck
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e
e
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r
en
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iv
etocon
i
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e
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asp
e
c
i
e
smoun
t
a
inch
i
ck
ad
e
e
sfo
r
ag
e(Sou
thwood
1961
).
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e
r
a
l
l
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efoundn
e
i
th
e
rth
eh
ab
i
t
a
tind
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rv
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e
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t
ioncompo
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iono
fn
e
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ts
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e
s
in
f
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c
edo
f
f
sp
r
ings
ex
.How
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enw
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e
s
t
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cu
r
r
inga
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ro
s
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r
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to
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u
rb
anh
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t
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eh
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l
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ex
.Inu
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r
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i
.
e
.
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c
r
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a
s
ing‘u
rb
an
n
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’
)
. Wh
i
l
eb
roods
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a
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io
sinth
eru
r
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h
ab
i
t
a
tv
a
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i
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ro
s
sth
es
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ea
stho
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einth
eu
rb
an
,th
e
r
ew
a
sl
i
t
t
l
etonov
a
r
i
a
t
ionin
48

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

th
eh
ab
i
t
a
tind
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c
e
so
fou
rru
r
a
ls
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t
e
s
.Thu
s
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e
r
ei
sl
i
t
t
l
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a
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t
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i
t
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r
ad
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sinou
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r
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ls
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t
e
sfo
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i
chf
em
a
l
emoun
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a
inch
i
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ad
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ldad
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t
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ju
s
t
o
f
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sp
r
ings
ex
.Bycomp
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r
i
son
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eh
ab
i
t
a
ta
tou
ru
rb
ans
i
t
e
sv
a
r
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r
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e
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t
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,
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r
ad
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aya
l
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r
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eg
i
cf
em
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l
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a
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rood
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e
r
e
l
a
t
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sh
ipb
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e
enh
ab
i
t
a
tu
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an
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z
a
t
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r
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a
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s
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t
etoou
rp
r
ed
i
c
t
ion
s
b
a
s
edonth
ef
ind
ing
so
fM
a
r
in
ie
ta
l
.(2017
)
.Thu
s
,howmoun
t
a
inch
i
ck
ad
e
e
sp
e
r
c
e
iv
eth
e
r
e
l
a
t
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equ
a
l
i
tyo
fu
rb
anv
s
.ru
r
a
lh
ab
i
t
a
t
,andth
em
a
l
e
sb
r
e
ed
ingw
i
th
inth
em
,m
ayb
e
comp
l
ex
.
P
r
ev
iou
swo
rkinb
l
a
ckc
app
edch
i
ck
ad
e
e
sfoundnoev
id
en
c
efo
rs
exa
l
lo
c
a
t
ion
,and
sugg
e
s
t
edth
ea
s
sump
t
ion
so
fs
exa
l
lo
c
a
t
ionth
eo
rym
ayno
th
o
ldinth
i
ssp
e
c
i
e
s(R
am
s
aye
t
a
l
.2003
)
.I
ti
spo
s
s
ib
l
eth
e
s
am
ei
st
ru
efo
rou
rs
tudypopu
l
a
t
iono
fmoun
t
a
inch
i
ck
ad
e
e
s
.Th
e
T
r
iv
e
r
sand W
i
l
l
a
rd(1973
)hypo
th
e
s
i
sa
s
sum
e
sth
a
to
f
f
sp
r
ingcond
i
t
ioni
sco
r
r
e
l
a
t
edto
m
a
t
e
rn
a
lcond
i
t
iondu
r
ingb
r
e
ed
ing
.How
ev
e
r
,T
r
iv
e
r
sand W
i
l
l
a
rd(1973
)sugg
e
s
tth
i
s
a
s
sump
t
ionm
ayo
n
lyb
et
ru
efo
rsp
e
c
i
e
sw
i
thsm
a
l
l
,p
r
ed
i
c
t
ab
l
eb
roods
i
z
e
s
.Th
emoun
t
a
in
ch
i
ck
ad
e
e
sinou
rs
tudypopu
l
a
t
ionh
av
el
a
rg
e
,h
igh
lyv
a
r
i
ab
l
eb
roods
i
z
e
s(
r
ang
e1to8
n
e
s
t
l
ing
s
)
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s
,th
i
sa
s
sump
t
ionm
ayno
tho
ld
.Th
eCh
a
rnovmod
e
l(1982
)a
s
sum
e
sth
e
r
ep
rodu
c
t
iv
ev
a
lu
eo
fson
sv
s
.d
augh
t
e
r
si
sr
e
l
a
t
edtop
a
t
e
rn
a
la
t
t
r
ibu
t
e
sth
a
td
i
f
f
e
r
en
t
i
a
l
ly
b
en
e
f
i
tth
es
ex
e
s
.B
a
s
edont
h
i
smod
e
l
,w
ep
r
ed
i
c
t
edm
a
l
econd
i
t
ionm
igh
th
av
eane
f
f
e
c
ton
n
e
s
t
l
ings
ex
.W
ea
l
sop
r
ed
i
c
t
edth
a
tex
t
r
a
p
a
i
ro
f
f
sp
r
ingm
igh
tb
emo
r
el
ik
e
lytob
em
a
l
e
,a
s
f
em
a
l
ech
i
ck
ad
e
e
sa
r
eknowntos
e
ekb
e
t
t
e
r
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a
l
i
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a
l
e
sa
sex
t
r
ap
a
i
rp
a
r
tn
e
r
s(O
t
t
e
re
ta
l
.
1998
).How
ev
e
r
,w
efoundnoe
f
f
e
c
tr
eg
a
rd
inge
i
th
e
ro
fth
e
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ep
r
ed
i
c
t
ion
s
,sugg
e
s
t
ingth
e
e
s
t
im
a
t
e
so
fcond
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t
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i
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e
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tin
f
lu
en
c
ef
em
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l
er
ep
rodu
c
t
iv
ed
e
c
i
s
ion
s
,o
r
m
ayno
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i
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e
r
en
t
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a
l
lyb
en
e
f
i
ton
es
exov
e
rth
eo
th
e
r
.

49

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

Bo
thmod
e
l
so
fs
exa
l
lo
c
a
t
ion(T
r
iv
e
r
sand W
i
l
l
a
rd1973
;Ch
a
rnov1982
)a
s
sum
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e
p
rodu
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ff
em
a
l
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f
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sp
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t
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econd
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.L
ik
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l
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s
,
f
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ech
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s
t
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i
shdom
in
an
c
eh
i
e
r
a
r
ch
i
e
s(R
am
s
ayandR
a
t
c
l
i
f
f
e2003
;G
r
av
a
e
ta
l
.2012
)and
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e
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r
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e
r
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i
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ank
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,andintu
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ff
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a
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a
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a
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Wh
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th
ab
i
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a
tm
ayb
eanimpo
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t
an
t co
r
r
e
l
a
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eo
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f
sp
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exinmoun
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a
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ad
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.Th
econ
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r
awf
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l
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ev
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sw
eh
av
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e
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ta
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s
e
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to
ffoodav
a
i
l
ab
i
l
i
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,ov
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r
a
l
l
b
r
e
ed
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au
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emo
r
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f
sp
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a
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einmo
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eu
rb
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i
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edh
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i
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a
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a
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sp
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rgood
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i
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ea
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so
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av
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fch
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t
im
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esu
c
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a
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Po
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t
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t
r
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l
eo
t
id
er
ep
e
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tm
i
c
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s
a
t
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esw
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l
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l
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b
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t
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rcond
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t
ionp
rodu
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.— Mo
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.

54

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e
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i
s (2017
)

3
.7
. Tab
l
e
s
T
ab
l
e3
.1
.Summ
a
ryo
fmoun
t
a
inch
i
ck
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e
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e2014
–2016b
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son
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ign
edr
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15
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E
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T
ab
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χ
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)
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t
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r
e 0
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G
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56

181
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37
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E
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T
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57

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3
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.

58

M
.S
c
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s
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s(2017
)

0
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)
.

59

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

CHAPTER4
:CONCLUS
IONSANDFUTUREDIRECT
IONS
V
a
r
i
a
t
ioninind
iv
idu
a
lcond
i
t
ionandh
ab
i
t
a
tqu
a
l
i
tyin
f
lu
en
c
er
ep
rodu
c
t
iv
es
t
r
a
t
eg
i
e
sin
num
e
rou
ssongb
i
rdsp
e
c
i
e
s(
fo
rr
ev
i
ew
ss
e
eSh
e
ldon1998
;G
r
i
f
f
i
the
ta
l
.2002
)
.Th
ego
a
lo
f
th
i
sth
e
s
i
sw
a
stog
a
inin
s
igh
tin
towh
e
th
e
rth
e
s
ef
a
c
to
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sin
f
lu
en
c
er
ep
rodu
c
t
iv
ed
e
c
i
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ion
sin
moun
t
a
inch
i
ck
ad
e
e
s
.Sp
e
c
i
f
i
c
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l
ly
,Ia
sk
edhowind
iv
idu
a
lcond
i
t
ionandh
ab
i
t
a
tu
rb
an
i
z
a
t
ion
in
f
lu
en
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eth
el
ik
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rodu
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eas
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i
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s
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rood
.I
foundev
id
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etosugg
e
s
tm
a
l
econd
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t
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t
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an
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ep
rodu
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s
ion
sinmoun
t
a
inch
i
ck
ad
e
e
s
.Ifoundm
a
l
e
,bu
tno
tf
em
a
l
e
,cond
i
t
ionin
f
lu
en
c
edEPP
,
w
i
tho
ld
e
rm
a
l
e
st
end
ingtoh
av
ealow
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rp
ropo
r
t
iono
fEPOinth
e
i
rn
e
s
t
s
.Inadd
i
t
ion
,
f
em
a
l
e
st
end
edtoeng
ag
einEP
C
sw
i
tho
ld
e
rm
a
l
e
s
.Th
e
s
ef
ind
ing
ssugg
e
s
tEPPi
sgov
e
rn
ed
byth
egoodg
en
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shypo
th
e
s
i
sinmoun
t
a
inch
i
ck
ad
e
e
s
,a
sh
a
sb
e
endo
cum
en
t
edino
th
e
r
P
a
r
id
a
esp
e
c
i
e
s(
e
.g
.
,K
emp
en
a
e
r
se
ta
l
.1992
;O
t
t
e
re
ta
l
.1998
).
P
r
ev
iou
swo
rkinmys
tudypopu
l
a
t
ionc
l
a
s
s
i
f
i
edmoun
t
a
inch
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ad
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sa
s‘u
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an
ad
ap
to
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s
,
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sth
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ssp
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c
i
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sab
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c
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s
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fu
l
lyinh
ab
i
t
,andapp
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a
r
stob
en
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f
i
tf
romr
e
s
id
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in
,u
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an
i
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edh
ab
i
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a
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a
r
in
ie
ta
l
.2017
)
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l
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ed
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edEPPtob
el
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s
sf
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equ
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sum
ab
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qu
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tyu
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ein
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ev
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r
ad
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t
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rb
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i
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ion
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eu
rb
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emo
r
e
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ik
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ef
em
a
l
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i
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ssu
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l
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ig
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l
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ab
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,EPP

60

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

r
a
t
e
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e
r
eh
igh
e
s
ta
tth
eh
ab
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a
tedg
eandin
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romth
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round
ingru
r
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)(Sm
i
the
ta
l
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l
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s
e
c
tabund
an
c
eandd
iv
e
r
s
i
ty(Sou
thwood1961
)
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s
equ
en
t
ly
,
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rb
anh
ab
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a
tm
ayb
emo
r
er
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e
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aZ
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r
t
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ta
l
.2017
;M
a
r
in
ie
ta
l
.2017
)
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c
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s
.B
e
c
au
s
emy
61

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

f
ind
ing
sr
eg
a
rd
ingh
ab
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t
a
tu
rb
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t
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in
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lythough
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. Wh
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.(2017
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4
.1
. Fu
tu
r
ed
i
r
e
c
t
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s
1
) Ino
rd
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r
th
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62

E
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i
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aS
.Bond
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rud

M
.S
c
.Th
e
s
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s (2017
)

2
) Ifoundp
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) F
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rex
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r
s
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tyhypo
th
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s
i
so
fEPP(G
r
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f
f
i
the
ta
l
.2002
)
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a
l
cu
l
a
t
ingth
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h
e
t
e
ro
zygo
s
i
tyo
fe
a
chind
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.

63

E
r
i
c
aS
.Bond
e
rud

M
.S
c
.Th
e
s
i
s (2017
)

4
.2
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c
lu
s
ion
s
Incon
c
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ry
. Mywo
rkh
a
s
exp
and
edonth
ef
ind
ing
so
fM
a
r
in
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ta
l
.(2017
)
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rov
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edn
ewin
s
igh
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toth
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ep
rodu
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s
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si
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lyon
emo
r
ep
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etoth
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tdyn
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4
.3
. L
i
t
e
ra
tu
r
ec
i
t
ed
B
l
a
i
r
,R
.B
.(
1996)
.L
andu
s
eandav
i
ansp
e
c
i
e
sd
iv
e
r
s
i
tya
longanu
rb
ang
r
ad
i
en
t
.
—E
co
l
.
App
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:506519
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G
r
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f
f
i
th
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,Ow
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an,K
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.(
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.Ex
t
r
ap
a
i
rp
a
t
e
rn
i
tyinb
i
rd
s
:ar
ev
i
ew
o
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t
e
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sp
e
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i
f
i
cv
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r
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ionandad
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iv
efun
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ion
.— Mo
l
.E
co
l
.11
:21952212
.
K
emp
en
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e
r
s
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.
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e
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en,G
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.
,V
and
enB
ro
e
ck, M
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,
Bu
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e,T
.
,V
anB
ro
e
ckhov
en
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t,A
.(
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.Ex
t
r
a
p
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i
rp
a
t
e
rn
i
tyr
e
su
l
t
sf
romf
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efo
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igh
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l
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lu
et
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t
.— N
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tu
r
e357
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.
L
aZ
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r
t
e
,S
.E
.
,O
t
t
e
r
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l
abb
ekoo
rn,H
.(2017
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. Moun
t
a
inch
i
ck
ad
e
e
sad
ju
s
tsong
s
,
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a
l
l
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s
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en
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im
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en
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c
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e
.— U
rb
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sy
s
t
.
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a
r
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.
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r,K
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aZ
e
r
t
e
,S
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ink, M
.W
.(
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rb
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i
ronm
en
t
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r
e
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s
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t
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th
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r
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a
tu
r
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l
h
ab
i
t
a
t
s
.— U
rb
anE
co
sy
s
t.(
Inp
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e
sp
r
e
f
e
rh
igh
r
ank
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a
l
e
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sex
t
r
a
p
ai
rp
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r
tn
e
r
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?— B
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,T
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.Th
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e
ro
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c
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e
so
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s
e
c
t
sa
s
so
c
i
a
t
edw
i
thv
a
r
iou
st
r
e
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s
.—
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T
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iv
e
r
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l
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.N
a
tu
r
a
ls
e
l
e
c
t
iono
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a
r
en
t
a
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i
l
i
tytov
a
ryth
es
ex
r
a
t
ioo
fo
f
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.— S
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65