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ge
co
s
y
s
t
em
s(K
e
an
ee
ta
l
.2
0
0
4
)
. To doso
e
f
f
e
c
t
i
v
e
l
yr
equ
i
r
e
sm
e
ch
an
i
s
t
i
cs
imu
l
a
t
ion
so
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l
im
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an
g
e
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and m
an
a
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co
lo
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lsu
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e
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s
ion
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d
i
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tu
rb
an
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tafin
esp
a
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i
a
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r
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en
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r
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th
th
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c
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t wh
i
chhum
and
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ak
in
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ak
e
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l
a
c
e
(Bon
ee
ta
l
.2
0
1
3
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e
an
ee
ta
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1
3
,K
in
ge
ta
l
.2
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1
3
)
.
2
2
6
Janua
ry2015
S
IMULATED CL
IMATEIMPACTS ON VEGETAT
ION
Dynam
i
cg
loba
lv
eg
e
ta
t
ion mod
e
l
s(DGVM
s
)a
r
ea
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ead
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eth
e po
t
en
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rchang
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e
ta
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tu
r
ec
l
ima
t
e
chang
es
c
ena
r
io
s(Ku
cha
r
ike
ta
l
.2006
, Gav
ine
ta
l
.2007
,L
i
t
t
e
l
le
ta
l
.20
10
)
.
Th
e
s
e mod
e
l
sa
r
ed
iv
e
r
s
e
, bu
tin g
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ra
lin
c
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iog
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i
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rban
c
em
e
ch
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i
sm
stos
imu
la
t
echang
e
sinv
eg
e
ta
t
ion(Moo
r
c
ro
f
t
2006
)
. DGVM
shav
ea
l
sob
e
enu
s
eda
tr
eg
iona
ls
ca
l
e
s
to p
rov
id
e pub
l
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r
iva
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eland manag
e
r
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e
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e
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rma
t
iononhowc
l
ima
t
ea
f
f
e
c
t
sv
eg
e
ta
t
ion(Ko
cae
t
a
l
.2006
, Rog
e
r
se
ta
l
.2011
)
.Tob
eg
loba
l
lyapp
l
i
cab
l
e
,
how
ev
e
r
, DGVM
s mu
s
ts
imp
l
i
fyth
e
i
rr
ep
r
e
s
en
ta
t
iono
f
v
eg
e
ta
t
ion andland
s
cap
e
s
. Mo
s
t DGVM
sd
e
s
c
r
ib
e
v
eg
e
ta
t
ioninb
roadp
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tfun
c
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iona
ltyp
e
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s
i
ognom
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roup
s
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rthan sp
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i
fy
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iv
idua
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i
e
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eg
e
ta
t
ioncommun
i
t
i
e
s(Ba
ch
e
l
e
te
t
a
l
.2001
)
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i
l
esom
e DGVM
s mayin
c
lud
e mana
g
e
m
en
t(A
lban
ie
ta
l
.2006
)
, manag
em
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ti
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ts
imu
la
t
ed
m
e
chan
i
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t
i
ca
l
lyba
s
edonhumand
e
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i
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ro
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e
s
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add
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, mo
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t DGVM
su
s
e aspa
t
ia
lg
ra
intha
ti
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r
s
e
r than th
e spa
t
ia
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ra
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t wh
i
chland
-u
s
e
d
e
c
i
s
ion
sa
r
e mad
e(Hu
r
t
te
ta
l
. 1998
)
. Su
c
c
e
s
s
fu
l
mod
e
l
ingp
ro
j
e
c
t
shav
ein
co
rpo
ra
t
edland
-manag
em
en
t
chang
e
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to c
l
ima
t
e
s
en
s
i
t
iv
ev
eg
e
ta
t
ion mod
e
l
ing
(S
ch
e
l
l
e
r and M
lad
eno
f
f 2005
,S
ch
e
l
l
e
re
ta
l
. 20
07
)
,
bu
tth
e
s
ep
ro
j
e
c
t
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r
el
im
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edbyhav
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r
idd
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ly
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r
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i
ca
l
ly
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e
en‘
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c
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rodu
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tg
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r
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ed bylo
ca
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ronm
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ta
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ion
s
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nonan
th
ropog
en
i
cd
i
s
tu
rban
c
e
s
,andhumaninflu
en
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e
s
.
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t
en
t tha
t DGVM
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im
i
t th
e dynam
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c
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ep
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imu
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eth
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r
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ly
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i
s
t
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ly
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r no
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ta
l
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e
i
r
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ro
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t
ion
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r
e un
l
ik
e
lyto p
r
ed
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c
tfu
tu
r
ev
eg
e
ta
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dynam
i
c
sa
c
cu
ra
t
e
ly
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r
th
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rmo
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, DGVM
s mu
s
t‘
‘
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in
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ra
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rbon and nu
t
r
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t poo
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s
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en p
r
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po
t
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ia
lv
eg
e
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i
c
sf
rom th
e sp
e
c
ifi
ed
c
l
ima
t
eandso
i
linpu
t
s
. Cu
r
r
en
tv
eg
e
ta
t
ion
,how
ev
e
r
,
i
so
f
t
ens
t
rong
lyd
e
t
e
rm
in
edbyth
el
ega
cyo
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r
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iou
s
land u
s
e andland cov
e
r
.P
r
io
r
i
ty e
f
f
e
c
t
s can b
e
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r
tan
td
e
t
e
rm
inan
t
so
ffu
tu
r
esu
c
c
e
s
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iona
ldynam
i
c
s(P
f
e
i
f
e
r
-M
e
i
s
t
e
re
ta
l
.2012
)
. Thu
s
, DGVM
s ma
y
p
rov
id
ep
ro
j
e
c
t
ion
stha
tfu
r
th
e
rd
iv
e
rg
ef
romcu
r
r
en
t
andfu
tu
r
econd
i
t
ion
si
fth
e
i
ra
s
s
e
s
sm
en
to
fg
row
ing
cond
i
t
ion
s do
e
s no
t ma
t
ch tha
to
f th
e cu
r
r
en
t
land
s
cap
e(Van Vuu
r
ene
ta
l
.2011
)
.
Fo
rth
e
s
er
ea
son
s
,th
r
e
ee
co
log
i
ca
l mod
e
l
inginno
v
a
t
ion
s wou
ldsub
s
tan
t
ia
l
ly advan
c
es
imu
la
t
ion
so
fc
l
i
ma
t
e
chang
e impa
c
t
s on v
eg
e
ta
t
ion in human
dom
ina
t
edland
s
cap
e
s
. Th
efi
r
s
ti
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e DGVM
p
ro
j
e
c
t
ion
sw
i
th mod
e
l
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ra
t
ea
tsu
ffi
c
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en
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l
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ra
in
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rban
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e
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e
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r
r
en
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eg
e
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t
iona
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r
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2
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t
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and m
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gh
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ingsy
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imu
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’
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eg
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ep
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econd
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r
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imu
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s
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eV
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imu
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emon
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ro
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and
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ap
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s
tu
rb
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e com
in
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en
tu
r
y
.
MATERIALS AND METHODS
M
o
d
e
ld
e
s
i
g
n
CV
-STSMin
t
e
g
r
a
t
e
sinpu
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ing f
ram
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i
g
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ak
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sav
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g
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t
a
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l
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i
th
inth
ea
g
en
t
-
2
28
GABR
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. YOSP
INET AL
.
1
1
b
a
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ed mod
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l Env
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va
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e
a(o
re
s
c
ap
ef
rom
ho
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t
i
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l
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r
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t
in
g
s
)tofo
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rco
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e
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t
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e
e sp
e
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i
e
sth
a
t cou
ld r
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en
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r
e MC
1 PVT
s
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ad
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el
and
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e
. W
eth
e
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fo
r
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th
a
tth
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g
r
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t
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e
r
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gdom
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an
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eo
fn
ew
t
r
e
esp
e
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i
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sw
a
s no
tn
e
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e
s
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r
ytos
imu
l
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t
e du
r
in
gth
e
r
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l
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ysho
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3y
e
a
r
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im
ef
r
am
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rs
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ed
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ed1
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i
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r
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es
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t
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e
s
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l
ink
ed e
a
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r typ
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r
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e
spond
in
g po
t
en
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ia
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e
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e
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a
t
iont
yp
e
sf
rom MC
1(T
ab
l
e1
)
. Tod
efin
es
t
a
t
e
s
,
w
eu
s
edfou
rp
a
r
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:dom
in
an
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i
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th
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t
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)
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)
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)
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)
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ad
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and
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ip
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edth
emto
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r
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e fou
rp
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r
sn
e
ed
ed to a
s
s
i
gn a
v
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t
ions
t
a
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etoe
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chIDU
.W
ee
sp
e
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i
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ys
c
ru
t
in
i
z
ed
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e
s
ed
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sto d
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aks
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h
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en
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.
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ran
s
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robab
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l
i
tyin
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l
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t
ion.—To g
en
e
ra
t
e
t
r
an
s
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t
ionp
rob
ab
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l
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t
i
e
sfo
rou
rSTSM
,w
eapp
l
i
edth
e
r
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e
so
ft
r
e
eg
row
thandsu
c
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lch
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g
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ro
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l
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v
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s
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fth
eP
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c
ifi
c No
r
thw
e
s
t
v
a
r
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an
to
f FVS(Gou
ld e
ta
l
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0
1
1
)
. Th
er
e
v
i
s
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s
in
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rpo
r
a
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ifi
c
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t
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stoth
eh
e
i
gh
t
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row
thr
a
t
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,
comp
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i
t
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v
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yn
am
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c
s
,and mo
r
t
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l
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yo
fO
r
e
gon wh
i
t
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o
akth
a
tw
e
r
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s
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e
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sp
a
r
to
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rl
a
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a
r
ch
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f
fo
r
t
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er
anFVSu
s
in
g mu
l
t
ip
l
et
r
e
el
i
s
t
stoin
i
t
i
a
l
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z
e
andt
r
a
inth
et
r
an
s
i
t
ionp
rob
ab
i
l
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t
i
e
sinou
rSTSM
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r
e
e
l
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s
t
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am
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romo
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lo
t
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roms
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e
r
a
ld
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t
sin
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e
rn and w
e
s
t
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rn O
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e
gon
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r
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c
ip
a
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romth
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r
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s
tIn
v
en
to
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yand An
a
l
y
s
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ab
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e(Woud
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r
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4
Th
et
r
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el
i
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t
sw
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r
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inn
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0
1
0
;a
v
a
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a
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l
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n
l
i
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e)
.
in
to STSM s
ta
t
e
s ba
s
ed on th
e fou
r pa
ram
e
t
e
r
s
d
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s
c
r
ib
edabo
v
e
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lo
td
a
t
af
rom w
e
s
t
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rn O
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gon(W
i
l
l
am
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eV
a
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ytosou
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r
r
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t
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. To
s
imu
l
a
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et
r
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e
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r
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eu
s
eda
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r
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rom
o
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ed
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in
gd
a
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a
. FVSrun
s
w
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r
ep
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r
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rm
edw
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thandw
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thou
tt
r
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r
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ane
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et
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sto d
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en
t andfu
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r
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and
s
c
ap
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s
,andth
enr
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-b
inn
edth
eou
tpu
tt
r
e
el
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s
t
sin
to
STSMs
t
a
t
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s
. Ou
ru
s
eo
f mu
l
t
ip
l
et
r
e
el
i
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t
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re
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chs
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a
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e
su
l
t
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an
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s
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lt
r
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rom wh
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ch w
ec
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l
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l
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t
edth
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a
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so
fannu
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s
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t
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rob
ab
i
l
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t
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s
.
1
4h
t
tp
:
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s
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f
ed
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s
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Janua
ry2015
S
IMULATED CL
IMATEIMPACTS ON VEGETAT
ION
2
3
1
FIG.2
. Sa
t
e
l
l
i
t
eim
a
g
e
r
yo
fth
es
tud
ya
r
e
alo
c
a
t
ioninth
eW
i
l
l
am
e
t
t
eV
a
l
l
e
y
,O
r
e
gon
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er
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e
c
t
an
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er
ep
r
e
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en
t
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0
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.
6
90
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bound
ing boxfo
rth
e MC
1s
imu
l
a
t
ion
s
. Th
eb
lu
el
in
esu
r
round
sth
es
tud
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a
. Sou
r
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e
: Goo
g
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th
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W
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122
8
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T
ran
s
i
t
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robab
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l
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tyad
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s
tm
en
t
sandcon
s
t
ra
in
t
s.—
1
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s
tom
i
za
t
ion—A ma
jo
rgoa
lo
fou
rp
ro
j
e
c
t
wa
stod
ev
e
lopa wa
ytob
e
g
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rs
imu
la
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sonth
e
p
r
e
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en
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s
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ap
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e
r
s
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f MC1th
a
t ma
t
ch
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s
e
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ed h
i
s
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i
c
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eg
e
ta
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, ra
th
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ro
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id
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im
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e
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t
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a
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e
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ion
. Do
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e
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l
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ene
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l
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1
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)
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r
ev
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ss
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la
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th MC1(Ro
g
e
r
se
ta
l
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1
1
)
,th
e
h
i
s
to
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i
c
a
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e
r
iod(1
8
95
–2
00
6
)y
i
e
ld
edat
emp
e
ra
t
em
a
r
i
t
im
ee
v
e
rg
r
e
en n
e
ed
l
e
l
e
a
ffo
r
e
s
t(wh
i
ch w
eequ
a
t
eto
Dou
g
la
s
-fi
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r
e
s
tinou
rs
tud
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)
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s
t
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ado
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e
p
r
a
i
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t
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t
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te
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th
inth
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y
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l
a
t
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en
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r
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ab
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ck1
9
6
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)
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ti
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e
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a
tth
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e
r
g
en
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r
eb
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e
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d
r
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t
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c
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l
l
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e
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e
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t
im
a
t
edh
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s
to
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r
ef
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equ
en
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,wh
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ch
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a
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en
c
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e
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r
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in
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g
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l
e
(Ch
r
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s
t
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l
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e
r
son2
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1
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xp
lo
r
eth
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s
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en
c
e
so
fth
i
s und
e
r
e
s
t
im
a
t
ion
,w
e bu
i
l
tn
ew ad
ju
s
t
ab
l
e
p
a
r
am
e
t
e
r
sin
to MC
1th
a
ta
l
low
edu
stosp
e
c
i
f
yth
efi
r
e
r
e
tu
rnin
t
e
r
v
a
l andth
e bu
rn d
a
t
e du
r
in
gth
esp
in
-up
ph
a
s
e
. W
eth
ene
x
am
in
ed MC
1ou
tpu
tv
a
r
i
ab
l
e
su
s
in
g
fi
r
er
e
tu
rnin
t
e
r
v
a
l
so
f3
,5
,1
0
,1
5
,and2
0y
e
a
r
s
,f
romth
e
1
5
0
thtoth
e3
0
0
thd
a
yo
fth
ey
e
a
r
,b
ys
t
ep
so
f2
5d
a
y
s
(
app
rop
r
i
a
t
efo
rth
efi
r
es
e
a
sono
fth
er
e
g
ion
’
sM
ed
i
t
e
r
r
an
e
anc
l
im
a
t
e
)du
r
in
gth
esp
in
-upph
a
s
e
.
TABLE 1
. S
ta
t
e
-and
t
r
an
s
i
t
ions
imu
l
a
t
ion mod
e
l(STSM
)t
r
e
eco
v
e
rt
yp
e
sw
i
thdom
in
an
tanda
s
so
c
i
a
t
edlo
c
a
lsp
e
c
i
e
s
.
STSMcov
e
rt
yp
e
Oaksavanna
Oak wood
land
Oakand Doug
la
s
-fi
r wood
l
and
Doug
la
s
-fi
randoak wood
l
and
Doug
la
s
-fi
rfo
r
e
s
t
Doug
la
s
-fi
rand map
l
efo
r
e
s
t
Doug
la
s
-fi
randg
randfi
rfo
r
e
s
t
B
ig
l
ea
f map
l
efo
r
e
s
t
P
in
esavanna
P
in
e wood
land
Mad
ron
e wood
landandfo
r
e
s
t
Mad
ron
eand Doug
l
a
s
-fi
rfo
r
e
s
t
Dom
in
an
tsp
e
c
i
e
s
d
e
s
c
r
ip
t
ion
Dom
in
an
tsp
e
c
i
e
s
d
rou
gh
t
to
l
e
r
an
tb
ro
ad
l
e
a
f
d
e
c
iduou
s
d
rou
gh
t
to
l
e
r
an
tb
ro
ad
l
e
a
f
d
e
c
iduou
s
d
rou
gh
t
to
l
e
r
an
tb
ro
ad
l
e
a
f
d
e
c
iduou
sandn
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
en
d
rou
gh
t
to
l
e
r
an
tb
ro
ad
l
e
a
f
d
e
c
iduou
sandn
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
en
l
e
s
sm
e
s
i
cn
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
en
m
e
s
i
cn
e
ed
l
e
l
e
a
fe
v
e
r
g
r
e
en
andb
ro
ad
l
e
a
fd
e
c
iduou
s
mo
r
em
e
s
i
cn
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
en
m
e
s
i
cb
ro
ad
l
e
a
fd
e
c
iduou
s
Q
u
e
r
c
u
sg
a
r
r
y
a
n
a
x
e
r
i
ce
v
e
r
g
r
e
enn
e
ed
l
e
l
e
a
f
x
e
r
i
ce
v
e
r
g
r
e
enn
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
enb
ro
ad
l
e
a
f
e
v
e
r
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32
GABR
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r
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ig
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imu
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en
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chan
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cfi
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l
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ap
(F
inn
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)
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s
ion(G
.I
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sp
in
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ep
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) wa
st
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r
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1
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;0
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andfo
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c
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a
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. A
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s show
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d
idno
ta
l
t
e
r MC
1
’
sp
ro
j
e
c
t
ion
so
fPVT
,andh
adon
l
y
m
ino
re
f
f
e
c
t
son MC
1ou
tpu
tv
a
r
i
ab
l
e
s
. Th
e
s
efind
in
g
s
ind
i
c
a
t
eth
a
ttu
rn
in
go
f
fth
efi
r
e mod
e
lin MC
1h
adon
l
y
am
ino
re
f
f
e
c
tonp
ro
j
e
c
t
ion
so
f MC
1ou
tpu
tv
a
r
i
ab
l
e
s
.
B
e
c
au
s
ew
etu
rn
edo
f
fth
efi
r
e mod
e
lin MC
1
,how
e
v
e
r
,
ou
rr
epo
r
t
ed r
e
su
l
t
s w
i
th MC1 w
i
l
ld
i
sa
g
r
e
e w
i
th
p
r
e
v
iou
sr
e
s
e
a
r
chu
s
in
g MC
1fo
rr
e
g
ion
sth
a
tin
c
lud
e
ou
rs
tud
ya
r
e
a(
e
.
g
.
, Ro
g
e
r
se
ta
l
.2
0
1
1
)
.
3
.T
r
a
n
s
i
t
i
o
np
r
o
b
a
b
i
l
i
t
ya
d
j
u
s
tm
e
n
t
s.—MC
1 ou
tpu
t
do
e
s no
tin
c
lud
eS
I
, wh
i
chi
sn
e
ed
ed b
y FVS
, bu
ti
t
in
c
lud
e
s num
e
rou
s ou
tpu
t
sr
e
l
a
t
edtov
e
g
e
t
a
t
ion p
ro
du
c
t
i
v
i
t
y
,so
i
l
s
, and c
l
im
a
t
e
. Tor
e
l
a
t
e MC
1 ou
tpu
t
v
a
r
i
ab
l
e
sto S
Iv
a
lu
e
s
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eco
r
r
e
l
a
t
ed a
v
e
r
a
g
ed MC
1
ou
tpu
tf
rom1
9
8
2
–
2
0
0
6w
i
tha
v
a
i
l
ab
l
eS
Id
a
t
a(Gou
lde
t
a
l
.2
0
1
1
)
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ep
e
r
fo
rm
edth
i
sco
r
r
e
l
a
t
ionfo
r1
7
9
0 MC
1
c
e
l
l
s
,r
an
g
in
gf
romc
en
t
r
a
ltosou
th
e
rn O
r
e
gon
,inth
e
Coa
s
t Rang
e and va
l
l
ey
s
. Th
i
sg
eog
raph
i
c cho
i
c
e
b
r
a
ck
e
t
sth
er
an
g
eo
fe
xp
e
c
t
ed S
Iinth
es
tud
ya
r
e
a
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romth
ep
r
e
s
en
tun
t
i
l2
0
9
9
,b
a
s
edonth
ep
r
ed
i
c
t
ionth
a
t
fu
tu
r
ec
l
im
a
t
einth
e W
i
l
l
am
e
t
t
eV
a
l
l
e
ym
a
yr
e
s
emb
l
e
cu
r
r
en
tc
l
im
a
t
einsou
thw
e
s
t
e
rn O
r
e
gon(Ro
g
e
r
se
ta
l
.
2
0
1
1
)
. W
eu
s
ed Ak
a
ik
e
’
sin
fo
rm
a
t
ionc
r
i
t
e
r
ion(A
IC
;
Ak
a
ik
e1
9
7
4
)tos
e
l
e
c
ta fin
a
l mod
e
lw
i
than o
v
e
r
a
l
l
2
¼0
.
5
5(T
ab
l
e2
)top
rodu
c
esp
a
c
e
-andt
im
e
ad
ju
s
t
edr
fi
l
l
in
gm
ap
so
fS
Ifo
rou
rs
tud
ya
r
e
a
.
Janua
ry2015
S
IMULATED CL
IMATEIMPACTS ON VEGETAT
ION
2
3
3
TABLE 2
. Va
r
iab
l
e
su
s
edinth
er
e
g
r
e
s
s
ionequ
a
t
ionr
e
l
a
t
in
g f
o
rth
et
r
an
s
i
t
ion
sth
a
tw
e
r
ed
i
s
a
l
low
edsoth
a
tth
esumo
f
MC1va
r
iab
l
e
stos
i
t
eind
e
x
.
a
l
low
edp
rob
ab
i
l
i
t
i
e
sw
a
sa
lw
a
y
sun
i
t
y
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sPVT
sch
an
g
e
Va
r
iab
l
e
Co
e
ffi
c
i
en
t
S
t
and
a
rd
i
z
ed
co
e
ffi
c
i
en
t
To
ta
lso
i
lca
rbon
9
.
0
8
13 1
04
Fo
r
e
s
tn
e
tp
r
ima
ryp
rodu
c
t
i
v
i
t
y 1
.
6
9
93 1
02
Av
e
rag
eJun
et
emp
e
r
a
tu
r
e
2
.
1
5
8
Abov
eg
roundl
iv
efo
r
e
s
tc
a
rbon 1
.
9
6
23 1
04
L
ea
fshap
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ex
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9
8
9
In
t
e
r
c
ep
t
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2
2
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1
8
3
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.
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5
5
1
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ea
fshap
eind
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an
g
e
scon
t
inuou
s
l
yf
rom0to1
,wh
e
r
e0
co
r
r
e
spond
sto n
e
ed
l
e
sh
ap
edl
e
a
v
e
s and 1 co
r
r
e
spond
sto
b
roadl
eav
e
s
.
Th
esp
a
t
i
a
lg
r
a
info
r MC
1(app
ro
x
im
a
t
e
ly80
0
-mg
r
id
c
e
l
l
s
)i
s mu
chla
rg
e
rth
anth
eIDU
s
,andthu
sth
e MC
1
p
ro
j
e
c
t
ed v
a
lu
e
sfo
rS
In
e
ed
edto b
e down
s
ca
l
edto
ind
i
v
idua
lIDU
s
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i
s
,w
ead
ju
s
t
edth
efu
tu
r
eS
Io
f
e
a
chIDU(S
I
)p
ropo
r
t
ion
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l
lyov
e
rt
im
e
,ba
s
edonth
a
t
IDU
t
)
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e MC
1c
e
l
l
’
s
in
i
t
ia
lS
I(S
I
,
IDU
’
s
in
i
t
ia
lS
I(S
I
IDU
0
MC
1
0)
andth
e MC
1c
e
l
l
’
sp
ro
j
e
c
t
edfu
tu
r
eS
I(S
I
)a
tt
im
et
MC
1t
IIDU0
S
IIDUt ¼S
S
IMC1t
:
S
IMC10
4
.T
ra
n
s
i
t
i
o
np
r
o
b
a
b
i
l
i
t
yc
o
n
s
t
ra
i
n
t
s.—S
in
c
eth
eSTSM
con
ta
in
sa
l
lv
e
g
e
ta
t
iontyp
e
sandt
r
an
s
i
t
ion
sn
e
ed
edfo
r
mu
l
t
ip
l
efu
tu
r
ec
l
im
a
t
es
c
ena
r
io
s
,a
sw
e
l
la
sth
ep
r
e
s
en
t
c
l
im
a
t
e
,w
el
im
i
t
edth
epo
s
s
ib
l
et
r
an
s
i
t
ion
sa
sfo
l
low
s
:(
1
)
w
ed
e
t
e
rm
in
ed wh
i
chco
v
e
rtyp
e
sanda
s
so
c
i
a
t
eds
t
a
t
e
s
andsp
e
c
i
e
s wou
ldb
ecomp
a
t
ib
l
ew
i
the
a
ch MC
1 PVT
and(2
)w
econ
s
t
ra
in
edth
eSTSMtoa
l
lowt
r
an
s
i
t
ion
s
on
lytocompa
t
ib
l
es
t
a
t
e
s(Tab
l
e3
)
.Fo
rex
amp
l
e
,und
e
r
th
esub
t
rop
i
c
a
lm
ix
edb
road
l
ea
fandn
e
ed
l
e
l
e
a
fev
e
r
g
r
e
en
fo
r
e
s
tPVT
,not
r
an
s
i
t
ion
stop
in
esa
vann
ao
r wood
l
and
a
r
ea
l
low
ed
,b
e
cau
s
e MC
1wou
ldno
tin
f
e
rth
eo
c
cu
r
r
en
c
e
o
fav
eg
e
t
a
t
iontyp
es
im
i
la
rtopond
e
ro
sap
in
esa
v
ann
ao
r
wood
l
andund
e
rth
i
sPVT
;fo
rth
esam
er
e
a
son
,e
x
i
s
t
in
g
p
in
e savanna o
r wood
land s
ta
t
e
s can on
ly r
em
a
in
un
ch
ang
ed
,o
rt
r
an
s
i
t
ionto o
th
e
rs
t
a
t
e
sv
ia a
l
low
ed
t
r
an
s
i
t
ion
s
.W
i
th
ine
a
chPVT
,t
r
an
s
i
t
ionp
rob
ab
i
l
i
t
i
e
sfo
r
e
a
chs
t
a
t
ew
e
r
ep
ropo
r
t
iona
l
l
yad
ju
s
t
edtocomp
en
s
a
t
e
inth
efu
tu
r
e
,w
ep
e
rm
i
ta
l
le
x
i
s
t
in
gs
t
a
t
e
stop
e
r
s
i
s
tinth
e
l
and
s
c
ap
e un
t
i
l ch
an
g
ed b
y su
c
c
e
s
s
ion o
r as
t
and
r
ep
l
a
c
in
gd
i
s
tu
rb
an
c
e
.I
ti
sc
l
e
a
rth
a
tc
l
im
a
t
ech
an
g
eh
a
s
in
c
r
e
a
s
edr
a
t
e
so
ft
r
e
e mo
r
t
a
l
i
t
yinsom
elo
c
a
t
ion
s(A
l
l
en
e
ta
l
.2
0
1
0
)
. MC
1ou
tpu
t
,how
e
v
e
r
,show
ednoe
v
id
en
c
e
o
fl
a
r
g
e
s
c
a
l
et
r
e
e mo
r
t
a
l
i
t
y(
e
.
g
.
,l
a
r
g
efu
tu
r
elo
s
s
e
so
f
t
r
e
ec
a
rbondu
etod
rou
gh
t
)
.B
e
c
au
s
eind
i
v
idu
a
lt
r
e
e
sa
r
e
lon
g
l
i
v
edandc
ana
c
c
l
im
a
t
eton
ewcond
i
t
ion
s
,th
e
yc
an
su
r
v
i
v
efo
rm
an
yy
e
a
r
sinsubop
t
im
a
lg
row
thcond
i
t
ion
s
,
l
e
ad
in
gtot
im
el
a
g
sandnon
l
in
e
a
rr
e
spon
s
e
stod
rou
gh
t
.
Fo
l
low
in
gas
t
and
r
ep
l
a
c
in
gd
i
s
tu
rb
an
c
e
,how
e
v
e
r
,on
l
y
t
r
an
s
i
t
ion
sa
l
low
edund
e
rth
ecu
r
r
en
tPVT m
a
yo
c
cu
r
.I
f
th
edom
in
an
tt
r
e
esp
e
c
i
e
sinth
ep
r
e
-d
i
s
tu
rb
an
c
es
t
a
t
ei
s
comp
a
t
ib
l
ew
i
thth
ecu
r
r
en
tPVT(T
ab
l
e3
)andi
sc
ap
ab
l
e
o
fr
e
sp
rou
t
in
gfo
l
low
in
gd
i
s
tu
rb
an
c
e(
i
.
e
.
,o
ak
s
,b
i
g
l
e
a
f
m
ap
l
e
,and m
ad
ron
e
)
,th
enth
eIDU w
i
l
lr
e
g
en
e
r
a
t
ea
sa
youn
gt
r
e
es
t
a
t
eo
fth
eco
v
e
rt
yp
eco
r
r
e
spond
in
gtoth
a
t
sp
e
c
i
e
s
.I
fth
ep
r
e
-d
i
s
tu
rb
an
c
es
t
a
t
ed
idno
tcon
t
a
inan
y
comp
a
t
ib
l
esp
e
c
i
e
sth
a
tr
e
sp
rou
tfo
l
low
in
gd
i
s
tu
rb
an
c
e
,
th
enth
eIDUr
e
g
en
e
r
a
t
e
sa
sanop
en
,g
r
a
s
s
/
fo
rbs
t
a
t
eo
f
th
e dom
in
an
tco
v
e
rt
yp
efo
rth
a
t PVT
. Fo
re
x
amp
l
e
,
w
i
th
inth
esub
t
rop
i
c
a
lm
i
x
ed b
ro
ad
l
e
a
fand n
e
ed
l
e
l
e
a
f
e
v
e
r
g
r
e
enfo
r
e
s
tPVT
,anIDUth
a
tw
a
sm
i
x
edo
akand
Dou
g
l
a
s
-fi
rwood
l
andp
r
e
-d
i
s
tu
rb
an
c
ew
i
l
lr
e
g
en
e
r
a
t
ea
sa
youn
go
aks
t
a
t
e
. Und
e
rth
a
ts
am
ePVT
,how
e
v
e
r
,anIDU
th
a
tw
a
sm
i
x
ed Dou
g
l
a
s
-fi
r and g
r
and fi
rfo
r
e
s
tw
i
l
l
r
e
g
en
e
r
a
t
ea
sa m
ad
ron
eg
r
a
s
s
/
fo
rb s
t
a
t
e andth
en
und
e
r
gop
rob
ab
i
l
i
s
t
i
csu
c
c
e
s
s
ion
.W
eadd
r
e
s
sth
el
im
i
t
a
t
ion
so
fth
i
sapp
ro
a
chinD
i
s
c
u
s
s
i
o
n.
MODEL BEHAVIOR AND RESULTS
STSMp
r
o
b
a
b
i
l
i
s
t
i
cb
e
h
a
v
i
o
r
F
i
r
s
t
,w
ep
e
r
fo
rm
ed(non
sp
a
t
i
a
l
)s
imu
l
a
t
ion
sou
t
s
id
e
o
f En
v
i
s
iontoe
x
am
in
eth
ep
rob
ab
i
l
i
s
t
i
cn
a
tu
r
eo
f CV
STSM
.S
t
a
r
t
in
gf
rome
i
th
e
rano
aks
a
v
ann
as
t
a
t
eo
ra
Dou
g
l
a
s
-fi
rs
t
a
t
e
,w
er
anr
ep
e
a
t
eds
imu
l
a
t
ion
sin CV
STSMfo
r5
0
0y
e
a
r
s
,w
i
th S
I¼ 3
0
.
4
8 m und
e
rth
e
m
a
r
i
t
im
ee
v
e
r
g
r
e
enn
e
ed
l
e
l
e
a
ffo
r
e
s
tPVT(F
i
g
.3
)
.W
e
TABLE 3
. S
ta
t
e
-and
t
r
an
s
i
t
ions
imu
l
a
t
ion mod
e
l(STSM
)t
r
e
eco
v
e
rt
yp
e
sto wh
i
cht
r
an
s
i
t
ion
sa
r
ea
l
low
edund
e
re
a
ch MC
1
d
e
r
iv
edpo
t
en
t
ia
lv
e
g
e
t
a
t
iont
yp
e(PVT
)
.
STSMcov
e
rt
yp
e
Oaksavanna
Oak wood
land
Oakand Doug
la
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2
34
GABR
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r mod
e
lrun
.A
s
CV
-STSMrun
sp
rob
ab
i
l
i
s
t
i
c
a
l
l
yfo
rlon
g
e
rp
e
r
iod
so
f
t
im
e
,lo
s
s
e
sb
e
g
intoa
c
c
ru
eino
ak wood
l
andon
c
eth
e
poo
lo
fo
aks
a
v
ann
ai
se
xh
au
s
t
ed(
e
.
g
.
,a
sinF
i
g
.3
)
,un
t
i
l
mo
s
to
f th
eland
s
cap
eb
e
com
e
s va
r
iou
s typ
e
so
f
Dou
g
l
a
s
-fi
rfo
r
e
s
t
,con
s
i
s
t
en
tw
i
thth
eth
eo
r
e
t
i
c
a
lc
l
i
m
a
t
i
c
c
l
im
a
xv
e
g
e
t
a
t
ionfo
rth
i
sa
r
e
a(F
r
ank
l
in and
D
y
rn
e
s
s1
9
8
8
)
. W
i
thsu
ffi
c
i
en
td
i
s
tu
rb
an
c
e
, how
e
v
e
r
,
th
e
r
ew
a
sas
t
ab
i
l
i
z
a
t
iono
rin
c
r
e
a
s
eino
aks
a
v
ann
aand
anin
c
r
e
a
s
einb
i
g
l
e
a
fm
ap
l
e
,a
c
comp
an
i
edb
yr
edu
c
ed
in
c
r
ea
s
e
sto Doug
la
s
-fi
rfo
r
e
s
t andlo
s
s
e
sto oak
wood
l
and
,con
s
i
s
t
en
tw
i
thth
eth
eo
r
e
t
i
c
a
lfi
r
ec
l
im
a
x
fo
rth
i
sa
r
e
a(H
ab
e
ck1
9
6
1
,F
r
ank
l
inand D
y
rn
e
s
s1
9
8
8
)
.
Th
ee
f
f
e
c
to
fc
l
im
a
t
ech
an
g
ew
a
sh
e
a
v
i
l
yd
ep
end
en
ton
d
i
s
tu
rban
c
e
, andin g
en
e
ra
l
,a
l
lc
l
ima
t
es
c
ena
r
io
s
p
rodu
c
eds
im
i
l
a
rch
an
g
e
sinv
e
g
e
t
a
t
ionfo
ran
yg
i
v
en
d
i
s
tu
rb
an
c
es
c
en
a
r
io b
yy
r9
0
,n
e
a
rth
e end o
fth
e
s
imu
l
a
t
ion(F
i
g
.4
)
. W
i
thou
td
i
s
tu
rb
an
c
e
,th
ec
l
im
a
t
e
ch
an
g
es
c
en
a
r
io
sh
adl
i
t
t
l
eimp
a
c
t on v
e
g
e
t
a
t
ion
.In
con
t
r
a
s
t
,und
e
rth
esho
r
t
e
s
td
i
s
tu
rb
an
c
er
e
tu
rnin
t
e
r
v
a
l
a
l
lo
fth
ec
l
im
a
t
e
ch
an
g
es
c
en
a
r
io
sg
en
e
r
a
t
ed al
a
r
g
e
r
edu
c
t
ioninth
eamoun
to
f Dou
g
l
a
s
-fi
rfo
r
e
s
tb
yth
eend
o
fth
e mod
e
lrunandin
c
r
e
a
s
edth
eamoun
to
f no
v
e
l
lo
c
a
lco
v
e
rt
yp
e
s(
e
.
g
.
,m
ad
ron
efo
r
e
s
t
)
.O
aks
a
v
ann
a
h
ab
i
t
a
t
sin
c
r
e
a
s
edw
i
thin
c
r
e
a
s
in
gd
i
s
tu
rb
an
c
er
e
g
a
rd
l
e
s
s
o
fp
ro
j
e
c
t
edc
l
im
a
t
ech
an
g
e and a
ch
i
e
v
edsom
ewh
a
t
h
i
gh
e
r qu
an
t
i
t
i
e
sth
an und
e
r no c
l
im
a
t
e ch
an
g
ea
t
i
s
tu
rb
an
c
el
e
v
e
l
s
.
comp
a
r
ab
l
ed
Th
ein
t
e
ra
c
t
ion b
e
tw
e
en d
i
s
tu
rb
an
c
e and c
l
im
a
t
e
ch
an
g
ea
l
soshow
edsom
eo
fth
enu
an
c
e
sd
i
f
f
e
r
en
t
i
a
t
in
g
howth
e GCM
sp
ro
j
e
c
tc
l
im
a
t
ech
an
g
efo
rth
es
tud
ya
r
e
a
(B
a
ch
e
l
e
te
ta
l
.2
0
1
1
, Ro
g
e
r
se
ta
l
.2
0
1
1
)
. Wh
i
l
ea
l
lth
r
e
e
GCM
sp
ro
j
e
c
t
edw
a
rm
in
ginth
es
tud
ya
r
e
a
,H
ad
l
e
yand
M
IROCp
ro
j
e
c
t
edsub
s
t
an
t
i
a
landcomp
a
r
ab
l
ew
a
rm
in
g
(
;4
.
28
C
)
, wh
i
l
e CS
IRO p
ro
j
e
c
t
edth
el
e
a
s
tw
a
rm
in
g
po
in
tint
im
e
.S
imu
l
a
t
ion
sw
e
r
econdu
c
t
ed w
i
thas
i
t
eind
e
xo
f3
0
.
4
8 mund
e
rth
em
a
r
i
t
im
ee
v
e
r
g
r
e
enn
e
ed
l
e
l
e
a
ffo
r
e
s
tpo
t
en
t
i
a
l
v
eg
e
ta
t
iontyp
e
.
2
36
E
co
log
i
ca
l App
l
i
ca
t
ion
s
Vo
l
.25
, No
.1
GABR
IELI
. YOSP
INET AL
.
(
;2
.6
8
C
)
. CS
IROp
ro
j
e
c
t
edala
rg
e
rin
c
r
ea
s
ein m
e
an
annua
lp
r
e
c
ip
i
ta
t
ion than M
IROC
, wh
i
l
e Had
l
ey
p
ro
j
e
c
t
edsma
l
ld
e
c
r
ea
s
e
sin m
eanannua
lp
r
e
c
ip
i
ta
t
ion
(Rog
e
r
se
ta
l
.2011
)
. Had
l
eyp
rodu
c
edth
e mo
s
tr
ap
id
chang
e
sinv
eg
e
ta
t
ion wh
end
i
s
tu
rban
c
eo
c
cu
r
r
edinth
e
l
and
s
cap
e
,w
i
thth
ela
rg
e
s
tin
c
r
ea
s
e
sinoakand mad
ron
e
cov
e
rtyp
e
s
. Byth
eend o
fth
es
imu
la
t
ion
s
, how
e
v
e
r
,
v
eg
e
ta
t
ion g
rown und
e
rth
eth
r
e
e GCM
st
end
edto
conv
e
rg
e
. M
IROCp
rodu
c
edl
i
t
t
l
echang
efo
r mo
s
to
f
th
es
imu
la
t
iont
im
e
.Th
era
t
eo
fv
eg
e
ta
t
ionchang
ea
tth
e
end o
f th
e M
IROC s
imu
la
t
ion
s w
i
th d
i
s
tu
rban
c
e
,
how
ev
e
r
,fa
rex
c
e
ed
edth
era
t
e
so
fchang
es
e
enund
e
r
th
eo
th
e
r comb
ina
t
ion
so
fc
l
ima
t
e and d
i
s
tu
rban
c
e
s
c
ena
r
io
s
. Pond
e
ro
sa p
in
e app
ea
r
ed
, bu
t on
ly v
e
ry
b
r
i
efly
,intwo ep
i
sod
e
s du
r
ingth
es
imu
la
t
ion(
s
e
e
App
end
ix C
)
.A
ti
t
sg
r
ea
t
e
s
tex
t
en
t(und
e
r Had
l
eyand
th
e17
%d
i
s
tu
rban
c
er
eg
im
e
)
,pond
e
ro
sap
in
ecov
e
rt
yp
e
s
o
c
cup
i
edapp
rox
ima
t
e
ly2
.8% o
fth
eIDU
sinth
es
tud
y
a
r
ea
.
Spa
t
ia
lou
tpu
t
G
iv
en th
e spa
t
ia
l
ly random and d
i
sagg
r
ega
t
ed
na
tu
r
eo
fth
ed
i
s
tu
rban
c
eu
s
edinth
e
s
es
imu
la
t
ion
s
,
i
ti
sinapp
rop
r
ia
t
etoov
e
r
in
t
e
rp
r
e
tth
espa
t
ia
lou
tpu
t
f
romth
ed
i
s
tu
rban
c
es
imu
la
t
ion
s
.W
i
thth
i
scav
ea
tin
m
ind
, how
ev
e
r
,th
espa
t
ia
l ou
tpu
tr
ev
ea
l
simpo
r
tan
t
mod
e
ldynam
i
c
s(F
ig
.5
)
.B
e
cau
s
eth
eimp
l
em
en
ta
t
ion
o
f CV
-STSMin Env
i
s
ioni
s ba
s
ed on t
ran
s
i
t
ion
p
robab
i
l
i
t
i
e
s op
e
ra
t
inginthou
sand
so
f IDU
s
,th
e
spa
t
ia
lou
tpu
tshowninF
ig
.5r
efl
e
c
t
sth
eou
t
com
e
so
f
thou
sand
so
find
ep
end
en
ts
imu
la
t
ion
s
. Th
ein
i
t
ia
l
land
s
cap
e con
ta
in
ed a m
ix
t
u
r
eo
f oak
, Doug
la
s
-fi
r
,
and b
ig
l
ea
f map
l
e cov
e
rtyp
e
s (F
ig
. 5A
)
. W
i
th a
d
i
s
tu
rban
c
er
e
tu
rnin
t
e
rva
lo
fs
ixy
r
sandnoc
l
ima
t
e
chang
e
, oak o
c
cu
r
r
edla
rg
e
ly onth
elow
e
rs
lop
e
so
f
th
e va
l
l
ey foo
th
i
l
l
s
, wh
i
l
e map
l
ela
rg
e
ly o
c
cu
r
r
ed
fu
r
th
e
r up
s
lop
e (F
ig
s
. 2 and 5B
)
. Con
t
empo
ra
ry
ob
s
e
rva
t
ion
s show a s
t
rong map
l
ep
r
e
s
en
c
e and
o
c
ca
s
iona
l dom
inan
c
ein up
land
sfo
l
low
ing d
i
s
tu
r
ban
c
e byt
imb
e
r ha
rv
e
s
t(
Johann
e
s
s
en e
ta
l
. 19
71
)
;
CV
-STSM ma
t
ch
e
sth
e
s
e ob
s
e
rva
t
ion
s
. Onth
eo
th
e
r
hand
,v
eg
e
ta
t
ion map
sf
rom1851dono
tshow map
l
e
dom
ina
t
ed s
tand
s in th
e up
land
s
,l
ik
e
ly b
e
cau
s
e
w
i
ldfi
r
e wa
sl
e
s
sf
r
equ
en
ta
th
igh
e
re
l
eva
t
ion
s away
f
romth
e va
l
l
ey floo
r
, andth
ec
l
ima
t
e wa
sl
ik
e
ly
som
ewha
tcoo
l
e
ra
tth
eendo
f‘
‘Th
eL
i
t
t
l
eI
c
e Ag
e
.
’
’
Th
es
t
rongin
t
e
ra
c
t
ion
sb
e
tw
e
enc
l
ima
t
echang
eand
d
i
s
tu
rban
c
ew
e
r
ea
l
soappa
r
en
tspa
t
ia
l
ly
.A
r
ea
so
fth
e
land
s
cap
etha
tex
i
s
t
eda
s Doug
la
s
-fi
rfo
r
e
s
tw
i
thou
ta
sub
s
tan
t
ia
loakcompon
en
ta
tth
es
ta
r
to
fth
es
imu
la
t
ion w
e
r
ea
lmo
s
ten
t
i
r
e
lyconv
e
r
t
edto mad
ron
eco
v
e
r
typ
e
s und
e
rth
eh
igh d
i
s
tu
rban
c
er
eg
im
e(F
ig
. 5C
)
.
Mu
ch o
f th
eland
s
cap
e
, aga
in
,e
sp
e
c
ia
l
lyin th
e
low
land
s
,r
ema
in
edino
rchang
edtoop
enoaksav
anna
cov
e
rtyp
e
s
. Th
i
s wa
s mo
s
tl
ik
e
ly du
etoth
e oak
’
s
ab
i
l
i
tytor
e
sp
rou
tfo
l
low
ings
tand
r
ep
la
c
ing d
i
s
tu
r
ban
c
e
.
DISCUSSION
Th
el
ik
e
l
ihoodth
a
t DGVM
sc
ana
c
cu
r
a
t
e
l
yp
r
ed
i
c
t
v
eg
e
t
a
t
ion dynam
i
c
s and d
i
s
tu
rban
c
einin
t
en
s
iv
e
ly
hum
an
-m
an
a
g
edl
and
s
c
ap
e
ssu
cha
sth
ew
i
ld
l
and
–u
rb
an
in
t
e
r
fa
c
ei
sr
emo
t
e
. DGVM
sa
r
e no
td
e
s
ign
ed to
m
e
ch
an
i
s
t
i
c
a
l
l
ys
imu
l
a
t
ean
th
ropo
g
en
i
cd
i
s
tu
rb
an
c
eo
r
r
e
a
l
i
s
t
i
cl
and
-u
s
ed
e
c
i
s
ion
-m
ak
in
ga
tafin
esp
a
t
i
a
lg
r
a
in
andl
a
ckt
im
el
a
g
scon
f
e
r
r
edb
yt
r
e
elon
g
e
v
i
t
y
.Inh
e
a
v
i
l
y
m
an
a
g
edl
and
s
c
ap
e
s
,fu
tu
r
es
imu
l
a
t
ion
sw
i
th DGVM
s
th
a
tp
ro
j
e
c
t po
t
en
t
i
a
ln
a
tu
r
a
lv
e
g
e
t
a
t
ion m
a
y no
tb
e
r
e
l
e
v
an
tto m
an
a
g
e
r
sin
t
e
r
e
s
t
edinch
an
g
e
stoe
x
i
s
t
in
g
l
andco
v
e
ro
v
e
rth
en
e
x
ts
e
v
e
r
a
ld
e
c
ad
e
s
.O
th
e
r mod
e
l
in
g
p
ro
j
e
c
t
s tha
t hav
ein
co
rpo
ra
t
ed manag
em
en
tin
to
DGVM
sa
r
eo
f
t
enun
ab
l
etod
e
l
i
v
e
rd
e
t
a
i
l
eda
s
s
e
s
sm
en
t
s
o
fl
and
co
v
e
rch
an
g
eo
v
e
rth
esho
r
tt
im
e ho
r
i
zon
sa
t
wh
i
chl
and
-u
s
ed
e
c
i
s
ion
sa
r
em
ad
e(S
ch
e
l
l
e
re
ta
l
.2
0
0
7
)
.
CV
-STSM
, th
e
r
e
fo
r
e
,p
rov
id
e
san
e
c
e
s
sa
ry b
r
idg
e
b
e
tw
e
en DGVM
s andl
and
-u
s
ed
e
c
i
s
ion mod
e
l
sth
a
t
w
i
shtos
imu
l
a
t
ech
an
g
e
stoe
x
i
s
t
in
gl
andco
v
e
r
.
CV
-STSM p
rodu
c
e
s exp
e
c
ted dynam
i
c
s w
i
th o
r
w
i
thou
td
i
s
tu
rb
an
c
e und
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e(K
ean
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a
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. 1996
, Ba
ch
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te
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. 2001
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ch
e
l
l
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ta
l
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0
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,
M
edv
igye
ta
l
.2009
, Mo
eu
rand Vand
end
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ch
e201
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)
.
How
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im
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ACKNOWLEDGMENTS
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LITERATURECITED
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, H
. 1974
. An
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