Flo a
208 (2013) 259–
267
Con en s
lis s
a ailable
a
SciVe se
ScienceDi ec
Flo a
jou
n
al
hom
epage:
www.else ie .com/loca e/ lo a
Long- e m
spa io- empo al
expansion
o
he
na i e-in asi e
Re ama
monospe ma
on
coas al
dunes:
Impo ance
o
land-use
and
na u al
dispe sal
ec o s
Sa a
Mu˜
noz-Vallés∗,
Juan
B.
Gallego-Fe nández,
Claudia
Dellafio e,
Jesús
Camb ollé
Depa amen o
de
Biología
Vege al
y
Ecología,
Uni e sidad
de
Se illa,
Ap.1095,
41080
Se ille,
Spain
a
i
c
l
e
i
n
o
A icle
his o y:
Recei ed
19
Janua y
2012
Recei ed
in
e ised
o m
3
Ma ch
2013
Accep ed
9
Ma ch
2013
A ailable online 1 May 2013
Keywo ds:
Dune
s abiliza ion
Land-co e
change
Seed
dispe sal
Li es ock
Wild
abbi
SW
Spain
a
b
s
a
c
Expansion
o
in asi e
species
inducing
sands
s abiliza ion
is
becoming
an
impo an
ecological
p oblem
in
coas al
dunes
in
some
pa s
o
he
wo ld.
Re ama
monospe ma,
an
endemic
o
coas s
o
SE
Spain
and
NE
o
Mo occo,
la e-colonizing
sh ub
occu ing
in
sandy
coas al
a eas,
was
plan ed
along
he
coas
o
SW
Spain
du ing
he
20 h
cen u y
o
s abilize
dunes.
In
ecen
decades,
he
species
has
sp ead
apidly,
becoming
in asi e,
and
some imes
incu ing
no able
changes
in
i s
en i onmen
and
plan
communi y.
The
expansion
pa e ns
o
R.
monospe ma
om
1956
o
2001
we e
desc ibed
and
quan ified
a
he
land-
scape
scale
wi hin
a
p o ec ed
dune
sys em,
using
GIS.
The
di e en ial
e ec s
on
expansion
pa e ns
o
he
main
ac o
con olling
he
popula ion
sp ead,
g azing
by
domes ic
li es ock
and
he
abundance
o
wild
abbi s,
we e
analyzed
compa a i ely.
R.
monospe ma
displays
an
exponen ial,
in asi e- ype
expansion
end,
p esen ing
a
mean
annual
co e age
inc ease
o
15%
and
a
mean
la e al
sp ead
a e
o
65.5
m
y −1
om
he
o iginal
popula ion
nucleus
o
he
Wes e n
and
Eas e n
ends
o
he
s udy
a ea.
The
dispe sal
ac i i y
p ima ily
o
abbi s
and
he
absence
o
compe ing
woody
species
con ibu e
o
his
apid
expan-
sion.
The
highes
local
inc eases
in
plan
co e age
could
be
ela ed
o
inc eased
abbi
abundance
and
o
he
imp o ed
ge mina ion
a es
o
seeds
which
ha e
passed
h ough
he
abbi
gu .
In
con as ,
a eas
highly
g azed
by
li es ock
p esen
he
lowes
inc eases,
and
an
open- ype
plan
communi y
is
main ained
he e
o
a
longe
ime.
© 2013 Else ie GmbH. All igh s ese ed.
In oduc ion
The
p ocess
o
coas al
dune
s abiliza ion
consis s
o
he
p og es-
si e
educ ion
o
sand
mo emen ,
closely
ela ed
o
an
inc ease
in
ege a ion
co e age,
and
ollowing
a
posi i e
eedback
mecha-
nism
(Yizhaq
e
al.,
2007).
The
p ocess
in ol es
significan
changes
in
dune
geomo phology
and
o he
en i onmen al
cha ac e is ics,
such
as
lowe
incidence
o
wind
and
sal
sp ay
and
inc eases
in
soil
o ganic
ma e
and
nu ien s
(El-Bana
e
al.,
2002;
Mu˜
noz
Vallés
e
al.,
2011)
as
well
as
in
he
ype
o
flo a
and
auna
sup-
po ed
by
he
dunes
(Lich e ,
1998;
Le in
and
Ben-Do ,
2004;
Plassmann
e
al.,
2010).
In
di e en
pa s
o
he
wo ld,
dune
s a-
biliza ion
is
ha ing
a
nega i e
impac
on
he
ichness
and
di e si y
o
dune
plan
and
animal
species
o
high
conse a ion
alue,
and
is
becoming
an
impo an
en i onmen al
p oblem
(A is,
1995;
Al a ez
and
Cushman,
2002;
Conse
and
Conno ,
2009;
Ise mann
e
al.,
2007;
Ise mann,
2008;
Ku iel
e
al.,
2004a;
Ma chan e
e
al.,
2008;
Wiedemann
and
Picka ,
1996).
A
p esen ,
fixa ion
and
∗Co esponding
au ho .
Tel.:
+34
954
557
165;
ax:
+34
954
626
308.
E-mail
add esses:
[email p o ec ed]
(S.
Mu˜
noz-Vallés),
[email p o ec ed]
(J.B.
Gallego-
Fe nández),
cdelafl[email p o ec ed]
(C.
Dellafio e),
[email p o ec ed]
(J.
Camb ollé).
s abiliza ion
o
coas al
dunes
a e
iden ified
by
se e al
au ho s
as
one
o
he
main
h ea s
o
hese
ecosys ems
(Ise mann
e
al.,
2007;
Jø gensen
and
Kollmann,
2008;
Ku iel
e
al.,
2004a).
The
ecen
inc ease
in
s abili y
o
coas al
dunes
has
been
caused,
in
ce ain
cases,
by
ac o s
such
as
global
o
local
clima ic
change
(Pana io
and
Pi˜
nei o,
1997;
Tsoa ,
2005;
Tsoa
e
al.,
2009)
and
changes
in
ag icul u al
and
li es ock
uses
(Ise mann
e
al.,
2007;
Le in
and
Ben-Do ,
2004;
Plassmann
e
al.,
2010);
o
as
a
esul
o
a
dec ease
in
he bi o y
by
domes ic
li es ock
o
wild
g aze s
such
as
abbi s
(Hodgkin,
1984;
Le in
and
Ben-Do ,
2004;
Ranwell,
1960,
1972;
Thomas,
1963).
In
many
cases,
howe e ,
dune
s abiliza ion
has
been
di ec ly
an h opogenic
and
is
he
p oduc
o
a emp s
o
p e en
dune
e osion
and
o
s abilize
blowing
ansg essi e
sand
shee s
(Pye,
1983).
Since
he
la e
18 h
cen u y,
a ious
coun ies
ha e
implemen ed
dune
fixa ion
and
s abiliza ion
echniques
ha
include
plan ing
apid-g ow h
he baceous
and/o
woody
species
ole an
o
en i onmen ally
s ess ul
dune
condi ions,
such
as
e.g.
Ammophila
a ena ia,
Hippophae
hamnoides,
Rosa
ugosa,
se e al
species
o
he
genus
Ca pob o us,
Acacia,
Eucalyp us,
Pinus
and
Re ama,
among
o he s,
wi h
no
ega d
o
hei
exo ic
cha ac e
in
e ms
o
he
ecosys em
in
which
hey
we e
plan ed
(A is,
1989;
Buell
e
al.,
1995;
Ise mann,
2008;
Ki h-y-Tassa a,
1946;
Kollmann
e
al.,
2009;
Ku iel
e
al.,
2004a,b;
Wiedemann
and
Picka ,
1996).
0367-2530/$
–
see
on
ma e ©
2013 Else ie GmbH. All igh s ese ed.
h p://dx.doi.o g/10.1016/j.flo a.2013.04.001
260 S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267
O e
ime,
many
o
hese
species
ha e
sp ead
beyond
hei
ini ial
plan a ion
a eas,
u he
s abilizing
he
su ounding
dune
sys ems,
and
some imes
exhibi ing
in asi e
beha iou
and
causing
se i-
ous
p oblems
o
he
conse a ion
o
he
coas al
dune
ecosys ems
(A is,
1995;
Conse
and
Conno ,
2009;
Ma chan e
e
al.,
2008;
Zunzunegui
e
al.,
2012).
Coas al
dune
a eas
a e
suscep ible
o
plan
in asion
because
o
he
open
na u e
o
he
plan
communi ies
and
exis ence
o
mic o-si es
sui able
o
species
es ablishmen ,
as
well
as
ypically
low
le els
o
plan –plan
compe i ion
(Jø gensen
and
Kollmann,
2008).
In
ecen
yea s,
he
impo ance
o
he
expansion
o
exo ic
species
in
coas al
dunes
and
hei
po en ial
o
induce
changes
in
he
s uc u e
and
unc ioning
o
esiden
communi ies,
ha e
been
ecognized
(Sakai
e
al.,
2001;
Valé y
e
al.,
2009).
A
limi ed
num-
be
o
s udies
deal
wi h
he
in asi e
expansion
o
na i e
species
wi hin
hei
na u al
a ea
o
dis ibu ion
(Pe e s
e
al.,
2006;
Van
Auken,
2000).
Such
s udies
a e
pa icula ly
sca ce
in
he
case
o
coas al
dunes
(Ise mann
e
al.,
2007;
Ise mann,
2008;
Nielsen
e
al.,
2011),
whe e
he
apid
expansion
and
inc ease
in
co e age
o
some
species,
pa icula ly
sh ubs,
significan ly
con ibu es
o
he
dune
s abiliza ion
p ocess,
al e ing
he
unc ioning
o
he
en i e
ecosys-
em.
An
analysis
o
his o ical
in asions,
sp ead
a es
and
pa e ns,
conduc ed
a
he
landscape
scale,
can
be
linked
o
p oxima e
causes
and
can
con ibu e
o
an
unde s anding
o
how
changes
in
s uc-
u e
and
unc ioning
o
dune
ecosys ems
occu
(Valé y
e
al.,
2009;
Vilà
and
Ibá˜
nez,
2011).
Such
analyses
can
also
be
used
o
model
u u e
sp ead
(Kollmann
e
al.,
2009),
as
well
as
help
o
de e mine
he
e ec i eness
o
di e en
managemen
p ac ices
(Ansley
e
al.,
2001).
Along
he
coas
o
SW
Spain,
Re ama
monospe ma
(L.)
Boiss.
(Tala e a,
1999),
a
N-fixing
leguminous
na i e
sh ub,
was
sown
in
coas al
dunes
du ing
he
20 h
cen u y,
ei he
alone
o
oge he
wi h
o he
species
like
Pinus
pinea
and
Ammophila
a ena ia,
in
o de
o
s abilize
he
dunes
(Ki h-y-Tassa a,
1946).
In
mos
o
he
loca-
ions
whe e
i
was
plan ed
wi hou
o he
species,
R.
monospe ma
has
displayed
in asi e
beha iou
and
has
apidly
expanded
beyond
he
plan a ion
a eas
(Valdés
e
al.,
2007).
I
has
caused
‘ e ama ’
o -
ma ions
in
high
densi ies
ha
ha e
led
o
sand
s abiliza ion
in
open,
semi-s abilized
dunes
and
induced
subs an ial
changes
in
dune
soil
and
plan
communi y,
as
well
as
a
he
landscape
le el
(Mu˜
noz
Vallés
e
al.,
2011;
Zunzunegui
e
al.,
2012).
In
addi ion,
he
species
is
p esen ly
conside ed
o
be
a
non-na i e
in asi e
plan
in
o he
coun ies
a ound
he
wo ld
such
as
USA
(Cali o nia:
Cal-IPC,
2006;
Randall,
1997;
Rejmanek
and
Randall,
1994)
and
Aus alia
(Randall,
2007).
The
expansion
o
a
plan
species
is
dependen
on
i s
li e
his-
o y
cha ac e is ics,
including
he
p oduc ion
and
dispe sal
o
p opagules,
he
cha ac e is ics
o
he
esiden
ecosys em
and
he
p e ailing
en i onmen al
condi ions
(Pe e s
e
al.,
2006).
In
his
ega d,
conse a ion
and
managemen
app oaches,
as
well
as
dis-
u bances,
can
a ec
popula ion
ends
in
ege a ion.
In
coas al
dunes,
wild
abbi s,
ha es
and
li es ock
a e
he
main
po en ial
dispe sal
agen s
o
R.
monospe ma
(Dellafio e
e
al.,
2006).
On
he
o he
hand,
he
adi ional
use
o
dunes
as
li es ock
g az-
ing
a eas,
o
he
p esence
o
wild
g aze s,
ha e
o en
con olled
he
excessi e
g ow h
o
plan
co e age
and
enc oachmen ,
becoming
an
impo an
ac o
in
he
main enance
o
low
ege a ion
co e age
in
semi-s abilized
dunes
and
hei
cha ac e is ic
open- ype
com-
muni ies
(Kooijman
and
Van
de
Meulen,
1996;
Le in
and
Ben-Do ,
2004;
Manzano
and
Ná a ,
2000;
Zhao
e
al.,
2005).
Land
use
ype
and
policies
o
na u e
conse a ion
and
managemen
could
he e-
o e
play
an
impo an
ole
in
con olling
ends
o
R.
monospe ma
expansion
in
he
coas al
dunes
whe e
he
species
g ows.
This
s udy
p esen s
a
de ailed
econs uc ion
o
he
his o i-
cal
expansion
o
R.
monospe ma
on
a
dune
landscape
by
using
ae ial
pho og aphy
and
GIS
analysis.
Specific
objec i es
we e:
(1)
o
model
he
expansion
o
R.
monospe ma
o e
he
dune
landscape,
(2)
o
find
ou
a ia ions
in
expansion
a es
and
spa io- empo al
ends,
and
(3)
o
es
how
he
p esence
o
li es ock
and
abun-
dance
o
wild
abbi s,
condi ioned
by
he
policies
o
na u al
a eas
conse a ion,
could
ac
o
influence
he
in asion
p ocess.
Ma e ials
and
me hods
S udy
a ea
The
s udy
was
conduc ed
on
he
El
Rompido
spi ,
a
semi-
isola ed
and
p o ec ed
dune
sys em
loca ed
in
he
Pied as
i e
es ua y,
in
he
cen al
coas al
egion
o
he
Gul
o
Cadiz,
SW
Spain
(37◦12N,
7◦07W),
and
included
in
he
na u al
ange
o
R.
monospe ma
(Fig.
1).
The
spi
s e ches
eas wa ds
o
app oxi-
ma ely
12
km
unning
pa allel
o
he
coas line,
is
be ween
300
and
700
m
in
wid h
and
cu en ly
co e s
an
a ea
o
534.7
ha
(Gallego-
Fe nández
e
al.,
2006).
The
spi
ac i ely
g ows
eas wa d
a
a
mean
annual
a e
o
a ound
42
m
o e
he
las
50
yea s,
adding
abou
1.5
ha
o
new
su ace
a ea
o
he
sys em
e e y
yea
(Mu˜
noz
Val-
lés,
unpub.
da a).
The
clima e
is
Medi e anean,
wi h
an
influence
om
he
A lan ic
Ocean.
Mean
annual
empe a u e
is
18.2 ◦C
and
mean
annual
ain all
is
583
mm,
wi h
a
p onounced
d ough
pe iod
om
May
o
Sep embe
(29
y .
eco d,
1974–2003;
Gib aleón
Me eo ological
S a ion,
Huel a).
The
spi
suppo s
di e en
eg-
e a ion
communi ies,
la gely
dependen
on
geomo phology
and
sedimen a y
ai s.
They
ange
om
high
beach
and
ac i e
dune
communi ies
o
in e io
semi-s abilized
and
s abilized
dunes,
in e -
dune
we
dep essions
o
‘slacks’,
sal
ma shes
and
idal
swales
(Gallego-Fe nández
e
al.,
2006).
A
he
beginning
o
he
wen ie h
Fig.
1.
The
El
Rompido
spi
s udy
a ea
on
he
coas
o
SW
Spain.
Limi s
o
P o ec ed
A ea,
g adien
o
age
o
R.
monospe ma
popula ion,
es ima ed
abundance
o
wild
abbi s
and
p essu e
o
g azing
by
li es ock
a e
indica ed.
The
loca ion
o
5
homogeneous
sec o s
selec ed
o
u he
compa isons
is
also
shown.
R.
monospe ma
co e age
on
he
spi
in
2001
is
ep esen ed
by
black
shading.
S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267 261
cen u y,
highly
mobile
sands
wi h
sca ce
ege a ion,
domina ed
by
he
psammophilous
Ammophila
a ena ia,
exis ed
in
he
a ea
(San
Miguel
de
la
Cáma a,
1913).
In
he
o ies
decade,
a
small
R.
monospe ma
popula ion
loca ed
in
he
cen al
sec o
o
he
spi
is
de ec able,
and
i
is
p esumable
ha
i
was
plan ed
he e
in
he
1920–30s
decades
(Gallego-Fe nández
e
al.,
2006).
A
p esen ,
he
species
is
widely
dis ibu ed
in
almos
all
he
inland
dunes.
Mos
o
he
spi
su ace
(73%)
has
been
included
in
he
p o ec ed
na u al
a ea
‘Pa aje
Na u al
Ma ismas
del
Río
Pied as
y
Flecha
del
Rompido’
since
1989
(Fig.
1),
and
in eg a ed
in o
he
Eu opean
NATURA
2000
ne wo k
(ES6150006).
Nine
habi a
ypes,
included
in
Annex
I
o
he
Eu opean
Habi a
Di ec i e
(Anonymous,
1992)
a e
ep esen ed
on
he
spi .
This
includes
semi-s abilized
dune
habi a s
(Eu opean
Communi ies
p o ec ed
habi a s
2130
and
2250)
whe e
R.
monospe ma
g ows.
In
addi ion,
ele en
endange ed
dune
species
and
subspecies
( wo
o
hem
p o ec ed
unde
law)
g owing
on
hese
dune
habi a s
ha e
been
eco ded
in
he
a ea
(Mu˜
noz
Vallés
e
al.,
2009).
The
spi
is
subjec
o
low
ou is
p essu e.
G azing
by
domes-
ic
li es ock
is
p ohibi ed
wi hin
he
p o ec ed
a ea;
ne e heless,
low
g azing
le els
s ill
occu
wi hin
a
educed
zone
close
o
he
limi s
o
he
p o ec ed
a ea.
Benefi ing
om
he
p o ec ion
poli-
cies
and
low
accessibili y
o
he
a ea,
an
impo an
popula ion
o
wild
abbi s
exis s
in
he
spi :
wa ens
a e
es ima ed
o
numbe
444
±
161
(mean
±
S.E.),
and
a e
mainly
concen a ed
in
he
cen al
and
cen al-eas e n
sec o s
(Dellafio e
e
al.,
2008)
–
Fig.
1.
S udy
species
Re ama
monospe ma
is
a
mid-successional,
N-fixing
woody
species
ound
in
coas al
sandy
a eas,
endemic
o
SW
Spain
and
NW
Mo occo.
Ne e heless,
i
success ully
es ablishes
in
o he
a eas
unde
Medi e anean
clima e
a ound
he
wo ld
(Mu˜
noz
Vallés
e
al.,
in
p ess).
Adul
plan s
a ain
heigh s
o
3–4.5
m,
canopy
diam-
e e s
o
up
o
10
m
and
indi idual
mean
co e age
o
up
o
70
m2.
Th ough
moni o ing
indi iduals
by
means
o
his o ical
ae ial
pho-
og aphy
(Mu˜
noz
Vallés
e
al.,
2005),
i
has
been
es ima ed
ha
a
li espan
o
55–80
yea s
is
possible.
Clonal
g ow h
has
no
been
documen ed
in
his
species,
howe e ,
i
can
p oduce
om
∼335
o
∼2800
ui s
pe
m2o
canopy
in
he
s udy
a ea,
om
la e
May
o
Sep embe .
The
ui s
lack
e iden
dispe sal
adap a ions,
and
me ely
all
o
he
g ound
whe e
hey
emain
a ailable
o
dispe se s
un il
he
nex
ui ing
season
(Mu˜
noz
Vallés
e
al.,
in
p ess).
The
apid
expansion
o
he
species
has
been
mainly
enhanced
by
he
dis-
pe sing
ac i i ies
o
he
wild
abbi
(O yc olagus
cuniculus),
as
well
as
an
imp o emen
in
seed
ge mina ion
a es
which
occu s
a e
he
passage
o
seeds
h ough
he
abbi
gu
( om
13%
o
24%;
Dellafio e
e
al.,
2006,
2010).
The
species
is
a
p esen
widely
dis ibu ed
in
he
s udy
a ea,
and
is
he
dominan
species
in
he
semi-s abilized
dunes,
which
ep esen
he
main
habi a
on
he
spi
(57%
o
he
o al
spi
su ace;
Gallego-Fe nández
e
al.,
2006).
Mapping
R.
monospe ma
co e age
This
s udy
was
based
upon
a
empo al
sequence
o
ae ial
o hopho og aphs,
co esponding
o
he
da es
1956
(B/W,
scale
1:33,000),
1977
(B/W,
scale
1:18,000),
1989
(colou ,
scale
1:5000)
and
2001
(colou ,
scale
1:5000).
Pho og aphs
we e
scanned
a
high
esolu ion
o
1200
dpi,
and
he
cen al
zone
o
each
one
was
selec ed
in
o de
o
a oid
edge
dis o ion.
They
we e
u he
geo- ec ified
using
G ound
Con ol
Poin s
aken
in
he
field
by
GPS.
Using
hese,
a
digi al
o hopho o
mosaic
was
c ea ed
o
each
da e
and
added
o
a
GIS
(A cView
3.2).
An
addi ional
ae ial
pho og aph
om
1946
(B/W,
scale
1:45,000)
was
examined.
I
was
no
added
o
he
GIS
due
o
low
image
quali y
bu
i
was
used
o
delimi
he
o iginal
popula ion
o
he
species
in
he
cen al
sec o
o
he
spi .
Indi idual
o
g oups
co e
o
R.
monospe ma
canopies
was
digi-
alized
in o
he
GIS
a
an
app oxima e
scale
o
1:500.
I
was
fi s
mapped
on
he
basis
o
he
mos
ecen
da e
mosaic
(2001),
wi h
he
highes
esolu ion
and
image
quali y,
and
subsequen ly
on
he
o hopho o
mosaics
om
he
da es
1989,
1977
and
1956.
The
wo k
on
each
da e
and
map
was
successi ely
suppo ed
in
he
p e i-
ously
buil
maps.
Due
o
he
a ailabili y
o
pho og aphs,
analysis
o
90%
o
he
cu en
o al
su ace
o
he
spi
was
possible.
This
excluded
a
sca ce
a ea
o
53.47
ha
loca ed
a
he
wes e n
ex eme.
Adul
indi iduals
o
R.
monospe ma
we e
easily
ecognizable
in
he
pho og aphs
due
o
hei
ex u e
and
one,
hei
cha ac e is-
ic
ci cula
o
ellipsoidal
shape
and
hei
size,
significan ly
la ge
han
adul s
o
he
o he
woody
species
p esen ,
such
as
A emisia
campes is
subsp.
c i hmi olia,
C ucianella
ma i ima
Helich ysum
i alicum
subsp.
pica dii
and
Thymus
ca nosus,
e en
om
a
young
age.
Analysis
o
o al
co e age
and
empo al
change
To al
analyzed
su ace,
semi-s abilized
dune
su ace
(sui able
o
he
es ablishmen
o
R.
monospe ma;
his
excludes
he
beach,
emb yo
dunes
and
sal
ma shes),
o al
co e age
o
he
species
and
pe cen age
o
co e age
pe
semi-s abilized
dune
su ace
we e
ob ained
om
he
GIS
o
e e y
s udied
da e.
Ra e
o
inc ease
in
co -
e age
was
subsequen ly
calcula ed
as
he
eg ession
o
he
squa e
oo
o
he
a ea
co e ed
by
he
species
as
a
unc ion
o
ime,
in
acco -
dance
wi h
p e ious
models
in
biological
in asion
heo y
(Buell
e
al.,
1995;
Skellam,
1951).
In
addi ion,
he
main
di ec ions
o
he
popula ions
la e al
sp ead
we e
iden ified
and
maximum
a es
o
la e al
sp ead
we e
subsequen ly
calcula ed
as
he
di e ences
in
he
popula ion’
limi s
be ween
da es.
Sp ead
owa ds
he
coas -
line
was
es ima ed
only
in
he
3
km
long
cen al
ame
o
he
spi ,
co esponding
o
he
o iginal
popula ion
nucleus.
Analysis
o
spa io- empo al
pa e ns
o
expansion
All
digi al
maps
o
dune
su ace
and
R.
monospe ma
co e age
we e
di ided
wi hin
he
GIS
in o
con iguous
50
m
×
50
m
cells
using
a
g id
(B own
and
Ca e ,
1998;
Middle on
e
al.,
2008).
Bo h
dune
su ace
and
co e age
o
R.
monospe ma
pe
dune
su ace
(%)
wi hin
each
cell
we e
calcula ed.
Due
o
he
he e ogenei y
o
he
a ea,
e.g.
dune
and
sal
ma shes,
only
hose
cells
in
which
>75%
o
he
su ace
was
dune
su ace
we e
conside ed
alid
cells,
and
we e
used
in
he
analyses.
This
app op ia e
cell
size
o
change
de ec-
ion
was
de e mined
a e
es ing
i
in
compa ison
wi h
ano he
wo
sizes
o
cells,
100
m
×
100
m
and
25
m
×
25
m.
The
amoun
o
100
m
×
100
m
cells
was
limi ed
by
he
he e ogenei y
o
he
a ea,
and
co e ed
a
smalle
su ace
han
he
50
m
×
50
m
ones
did.
On
he
o he
hand,
25
m
×
25
m
cells
we e
oo
small
in
size
and
apidly
eached
high
pe cen ages
o
R.
monospe ma
co e age.
Fo
hese
easons,
analyses
we e
finally
ca ied
ou
using
50
m
×
50
m
cells
only.
Cells
we e
classified
in o
fi e
ca ego ies
acco ding
o
he
co -
e age
o
R.
monospe ma
pe
dune
su ace,
as
ollows:
C0:
co .
=
0%;
C1:
0%
<
co .
≤
25%;
C2:
25%
<
co .
≤
50%;
C3:
50%
<
co .
≤
75%;
C4:
75%
<
co .
≤
100%.
The
o al
numbe
o
alid
cells
pe
ca ego y
and
da e
was
calcula ed,
and
maps
o
he
dis ibu ion
o
cell
ca ego ies
on
each
da e
we e
p oduced.
Analysis
o
spa io- empo al
change
in
ela ion
o
g azing
by
domes ic
li es ock
and
wild
abbi
abundance
Fi e
homogeneous
sec o s
o
1
km
in
leng h
each
we e
selec ed
h oughou
he
s udy
a ea,
acco ding
o
li es ock
g azing
in en-
si y
and
abundance
o
wild
abbi .
Cha ac e is ics
o
hese
sec o s
a e
indica ed
in
Table
1,
and
hei
loca ion
is
shown
in
Fig.
1.
262 S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267
Table
1
Cha ac e is ics
o
sec o s
(S1–S5)
selec ed
o
analysis
o
spa io- empo al
change
in
ela ion
o
land
use
and
conse a ion
le el.
S1
S2
S3
S4
S5
Included
in
he
p o ec ed
a ea
No
Yes
Yes
Yes
Yes
In ensi y
o
li es ock
g azingaHigh
Low
None
None
None
Abundance
o
wild
abbi bVe y
low(0.0)
Low(1.3)
High(2.1)
The
highes (3.0)
Ve y
low(0.5)
Dis ance
om
he
oldes
popula ion
nucleous
(km)c5.3
2.4
0
2.9
5.2
aAcco ding
o
Zunzunegui
e
al.
(2012).
bMean
numbe
o
wa ens
pe
ha,
acco ding
o
Dellafio e
e
al.
(2008).
cMeasu ed
om
he
cen e
o
he
sec o s.
Table
2
Desc ip i e
da a
abou
semi-s abilized
dune
a ea,
R.
monospe ma
co e ing
and
alid
cells
used
o
u he
analyses,
om
he
analysis
on
GIS
on
ou
s udy
da es
be ween
1956
and
2001.
1956
1977
1989
2001
Semi-s abilized
dune
su ace
(ha)
271.53
288.94
295.51
296.84
R.
monospe ma
o al
co e age
(ha)
15.03
44.24
75.10
116.19
R.
monospe ma
co e age
pe
semi-s abilized
dune
su ace
(%)
5.53
15.31
25.41
39.14
Numbe
o
alid
cells
(50
m
×
50
m) 823
877
882
884
Semi-s abilized
dune
su ace
included
in
he
alid
cells
(%)
68.95
71.80
71.40
71.22
R.
monospe ma
co e age
included
in
he
alid
cells
(%)
66.99
67.19
69.01
76.99
Li es ock
g azing
in ensi y
was
es ima ed
as
mean
numbe
o
d oppings
pe
m2and
mean
numbe
o
ui s
pe
m2 ound
below
R.
monospe ma
canopies,
acco ding
o
Zunzunegui
e
al.
(2012).
High
in ensi y
co esponded
o
app oxima ely
16
d oppings
and
less
han
5
ui s
pe
m2;
low
in ensi y
co esponded
o
app oxima ely
6
d oppings
and
167
ui s
pe
m2;
non-in ensi y
co esponded
o
0
d oppings
and
app oxima ely
1158
ui s
pe
m2.
Abundance
o
wild
abbi s
was
es ima ed
as
wa en
densi y,
acco ding
o
Dellafio e
e
al.
(2008)
(Table
1).
Fo
each
sec o ,
R.
monospe ma
co e age
(%
o
sui able
su ace)
pe
use ul
cell
and
da e
was
calcula ed.
Mean
co e age
pe
cell
was
compa ed
in
he
fi e
sec o s
o
each
da e
using
one-way
ANOVA
and
he
pos
hoc
Tukey
es .
The
a e
o
inc ease
in
co -
e age
was
calcula ed
wi hin
each
sec o ,
o e
he
cou se
o
he
45
yea s
s udied,
as
he
eg ession
o
he
squa e
oo
o
he
a ea
co e ed
by
he
species
as
a
unc ion
o
ime
(Buell
e
al.,
1995;
Skellam,
1951).
Da a
we e
ans o med
(a csine
x)
p io
o
analysis
in
o de
o
comply
wi h
he
assump ion
o
homogenei y.
Fo
s a is-
ical
analysis,
he
s a is ical
so wa e
package
STATISTICA
6.0
was
used.
Resul s
To al
co e age
o
R.
monospe ma
and
empo al
change
Due
o
g ow h
dynamics
o
he
spi ,
bo h
he
a ea
o
he
spi
in
o al
as
well
as
he
a ea
o
dunes
inc eased
o e
ime
(Table
2,
Fig.
2).
The
o al
a ea
co e ed
by
R.
monospe ma
subs an ially
inc eased
du ing
he
45
yea s
analyzed;
om
15.0
ha
occupied
in
1956
(5.5%
o
dune
su ace)
o
116.2
ha
(39.1%
o
dune
su ace)
in
2001.
Assum-
ing
a
linea
inc ease,
he e
was
an
annual
mean
inc ease
in
he
a ea
co e ed
by
he
species
o
almos
15%.
The
mean
annual
inc ease
in
co e age
was
highe
be ween
1956
and
1977
(9.3%),
bu
co e -
age
inc ease
in
ela ion
o
popula ion
size
was
lowe
be ween
1977
and
1989
(5.8%),
and
be ween
1989
and
2001
(4.6%):
Table
3.
The
annual
a e
o
inc ease
in
o al
sh ub
co e age
was
ound
o
fi
an
exponen ial
cu e
(Fig.
3).
Re ama
monospe ma
eached
he
wes e n
limi
o
he
digi al
maps
du ing
he
fi s
pe iod
(1956–1977),
and
eached
he
wes e n
end
o
he
spi
in
1989.
Wes wa d
la e al
sp ead
was
he
mos
apid
du ing
he
fi s
s udied
pe iod,
ad ancing
mo e
han
120
m
y −1
Fig.
2.
Dis ibu ion
o
R.
monospe ma
co e age
on
each
o
ou
di e en
da es
on
he
El
Rompido
spi ,
SW
Spain.
Abo e,
he
loca ion
o
he
o iginal
popula ion
co e
is
ma ked
by
discon inuous
line
o e
an
ae ial
pho og aph
om
1946.
S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267 263
Table
3
Ra es
o
R.
monospe ma
co e age
inc ease
and
la e al
sp ead
o
he
popula ion
limi s
in
each
o
he
h ee
dis inguished
pe iods
and
o e
he
o al
o
45
yea s.
Since
he
popula ion
eached
he
wes e n
end
o
he
spi
a
some
ime
be ween
1977
and
1989,
wes wa d
la e al
sp ead
du ing
he
pe iod
1977–1989
was
no
calcula ed,
bu
his
ad ance
was
ce ainly
inexis en
be ween
1989
and
2001.
1956–1977
1977–1989
1989–2001
1956–2001
To al
inc ease
o
R.
monospe ma
co e age
(%)
194.45
69.74
54.71
673.23
Annual
a e
o
inc ease
in
R.
monospe ma
co e age
(%)
9.26
5.81
4.56
14.96
Annual
a e
o
wes wa d
la e al
sp ead
(m
y −1)
55.76
77.17
17.00
51.13
Annual
a e
o
eas wa d
la e al
sp ead
(m
y −1)
>120.00
–
0.00
79.78
Annual
a e
o
la e al
sp ead
owa ds
he
coas line
(m
y −1)
1.57
−1.00
1.50
0.87
Fig.
3.
Inc ease
in
R.
monospe ma
co e age
on
he
El
Rompido
spi ,
SW
Spain,
as
unc ion
o
ime.
(Table
3).
In
his
case,
we
we e
no
able
o
calcula e
he
wes wa ds
ad ance
o
he
popula ion
du ing
he
pe iod
1977–1989,
bu
we
can
say
ha
his
ad ance
was
null
be ween
1989
and
2001
(Table
3).
The
eas wa d
ad ance
was
slowe
in
compa ison
wi h
he
wes wa ds
ad ance.
A
maximum
speed
o
77.2
m
y −1was
achie ed
be ween
1977
and
1989,
and
dec eased
a e wa ds
(Table
3).
By
1977,
he
popula ion
had
eached
an
a ea
e y
close
o
he
eas e n
ex eme
o
he
spi .
Ne e heless,
he
eas e n
end
o
he
spi
had
s ill
no
been
eached
by
2001
(Fig.
2).
La e al
sp ead
owa ds
he
coas line
was
be ween
one
and
wo
o de s
o
magni ude
slowe
han
sp ead
along
he
leng h
o
he
spi ,
a
a ound
0.9
m
y −1be ween
1956
and
2001,
and
in
ac
showed
a
e ea
be ween
1977
and
1989
(Table
3).
Pa e ns
o
R.
monospe ma
expansion
The
spi
g ow h
dynamics
p oduced
an
inc ease
in
he
numbe
o
alid
cells
om
823
o
884
o e
he
s udied
pe iod.
Valid
cells
Fig.
5.
Rela i e
abundance
o
alid
50
m
×
50
m
cells
belonging
o
each
o
fi e
co -
e age
ca ego ies,
as
pe cen age
o
he
o al
numbe
o
alid
cells,
on
each
o
ou
analyzed
da es.
co e ed
be ween
69%
and
72%
o
he
semi-s abilized
dune
su -
ace
and
be ween
67%
and
77%
o
he
digi alized
co e age
o
R.
monospe ma.
Ca ego ized
alues
o
R.
monospe ma
co e age
we e
in
2001
highe
a
he
inne ,
olde
colonized
cells
(co e age
ca ego ies
C3
and
C4),
which
we e
loca ed
in
he
cen al
a ea
o
he
spi ,
and
lowe
(C0
and
C1)
in
he
mo e
ecen ly
colonized
cells,
which
we e
loca ed
mo e
o
he
ex e io
pa s
o
he
spi
(Fig.
4).
The
pe cen age
o
emp y
cells
(C0)
was
highe
a
he
fi s
da e
analyzed,
1956,
bu
apidly
dec eased
o e
ime
as
he
popula ion
expanded
(Fig.
5).
F om
1956
o
1977,
he
popula ion
p e e en ially
expanded
eas wa d,
and
gene ally
main ained
a
low
co e age
(below
25%)
ou side
he
cen al
nucleus
a ea.
A
e e y
da e,
apa
om
1956,
he
C1
cells,
associa ed
wi h
coloniza ion
e en s,
we e
ound
o
be
he
mos
abundan
co e age
ca ego y.
The
pe cen age
o
C1
cells
inc eased
un il
1989,
when
C0
we e
below
8%,
and
dec eased
he e-
a e .
This
indica es
a
change
om
a
p edominan ly
coloniza ion
o
new
cells
o
a
pa e n
in
he
final
pe iod
when
he
popula ion
became
mo e
dense.
Cells
om
ca ego ies
C2,
C3
and
C4
we e
gene ally
less
abundan
han
C0
o
C1,
and
ollowed
a
gene al
inc easing
end
o e
ime
(Fig.
5).
The
cen al
nucleus
a ea
showed
mode a e
o
null
inc eases
in
co e age
om
1977
o
2001,
while
co e age
losses
we e
egis e ed
in
somewha
mo e
han
1%
o
cells
Fig.
4.
Dis ibu ion
o
ca ego ies
o
R.
monospe ma
co e age
(C0–C4)
in
50
m
×
50
m
cells
in
1956
(A)
and
2001
(B).

264 S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267
Fig.
6.
Inc eases
in
R.
monospe ma
co e age
in
5
selec ed
sec o s
o
he
El
Rompido
spi ,
on
each
o
ou
analyzed
da es.
S1:
ou side
he
p o ec ed
a ea,
high
in ensi y
o
li es ock
g azing,
e y
low
abbi
abundance;
S2:
p o ec ed,
low
in ensi y
o
li e-
s ock
g azing,
low
abbi
abundance;
S3:
p o ec ed,
no
g azing
by
li es ock,
high
abbi
abundance;
S4:
p o ec ed,
no
g azing
by
li es ock,
he
highes
abundance
o
abbi s;
S5:
p o ec ed,
no
g azing
by
li es ock,
e y
low
abbi
abundance.
As e isks
indica e
significan
di e ences
ound
by
ANOVA;
di e en
le e s
indica e
significan
di e ences.
om
1977
o
1989
and
1.8%
om
1989
o
2001.
O e
ime,
al hough
R.
monospe ma
colonized
wes wa d
mo e
quickly
han
eas wa d,
he
occu ences
became
dense
owa ds
he
eas .
This
pa e n
can
be
ela ed
wi h
he
di e en ial
e ec
o
g azing
by
domes ic
li e-
s ock
and
wild
abbi s
(Fig.
4).
Pa e ns
o
R.
monospe ma
expansion
in
ela ion
o
g azing
by
domes ic
li es ock
and
wild
abbi
abundance
G id
cells
co esponding
o
he
C1
co e age
ca ego y,
associa ed
wi h
coloniza ion
e en s,
inc eased
un il
1977
in
sec o s
S2
and
S4,
and
un il
1989
in
S5,
and
dec eased
a e wa ds.
The
maximum
numbe
o
C1
cells
in
S3
was
eached
in
1956,
dec easing
a e -
wa ds,
while
in
S1
he
maximum
was
eached
in
2001
(da a
no
p esen ed).
This
indica es
a
change
in
dynamics,
mainly
om
col-
oniza ion
o
new
cells
(majo i y
o
C1)
o
popula ion
densifica ion
(C1
dec ease),
which
a ied
be ween
sec o s
s udied.
ANOVA
e ealed
significan ly
highe
mean
alues
o
R.
monospe ma
co e age
pe
cell
in
S3,
he
popula ion
co e,
du -
ing
1959
and
1977
(1956:
F
=
65.10;
P
<
0.001;
1977:
F
=
71.64;
P
<
0.001),
while
he
emaining
sec o s
main ained
simila
and
lowe
alues
(Fig.
6).
Mean
co e age
pe
cell
con inued
o
be
sig-
nifican ly
highe
in
S3
in
1989,
while
alues
in
S4
significan ly
inc eased
in
compa ison
wi h
S1,
S2
and
S5
(F
=
59.73;
P
<
0.001).
In
2001,
mean
co e age
pe
cell
showed
significan ly
di e en
alues
in
e e y
sec o
(F
=
144.34;
P
<
0.001).
In
his
yea ,
he
highes
co -
e age
alues
we e
ob ained
in
S4,
exceeding
hose
o
he
popula ion
co e
(S3),
while
p og essi ely
lowe
alues
we e
ob ained
in
S3,
S2,
S5
and
S1
(Fig.
6).
The
annual
a e
o
inc ease
in
co e age
fi ed
an
exponen ial
cu e
in
sec o s
S1
(P
=
0.002;
R2=
0.99),
S2
(P
=
0.004;
R2=
0.99),
S4
(P
=
0.016;
R2=
0.97)
and
S5
(P
=
0.023;
R2=
0.96),
bu
i
fi ed
a
lineal
cu e
in
sec o
S3
(P
=
0.007;
R2=
0.99).
Discussion
Ou
s udy
shows
ha
Re ama
monospe ma
has
apidly
expanded
in
he
open,
semi-s abilized
dunes
o e
he
45
yea
pe iod
co -
e ed
by
he
da a,
om
an
o iginal
cen al
nucleus
es ima ed
o
ha e
been
ounded
in
he
1920s
o
1930s
(Gallego-Fe nández
e
al.,
2006).
Speed
o
la e al
sp ead
exceeded
108
m
y −1,
and
s ill
he
popula ion
is
ac i ely
colonizing
dune
su aces
ha
newly
become
a ailable.
A
subs an ial
inc ease
in
co e age
o
673%
occu ed
du -
ing
hose
45
yea s,
and
a
p esen
a
p edominan
dynamic
o
popula ion
densifica ion
can
be
egis e ed.
The
s and
ha e
he
po en ial
o
u he
enc oachmen .
This
expansion
has
in
u n
con ibu ed
o
he
s abiliza ion
o
much
o
he
dune
sys em,
induc-
ing
subs an ial
changes
in
he
dune
flo a
and
ege a ion
s uc u e
(Mu˜
noz
Vallés
e
al.,
2011;
Zunzunegui
e
al.,
2012).
Acco ding
o
Alpe
e
al.
(2000),
in
ecosys ems
cha ac e ized
by
s ess ul
en i onmen al
condi ions,
ce ain
egionally
na i e
species
may
display
mo e
acu ely
in asi e
expansion
as
compa ed
o
alien
species,
as
a
esul
o
being
be e
adap ed
o
hose
condi-
ions.
The
annual
g ow h
a e
in
co e age
o
R.
monospe ma
(15%)
was
significan ly
highe
han
he
2.9%
ound
o
he
alien
Aca-
cia
saligna
in
an
analysis
o
da a
o e
34
yea s
by
Ku iel
e
al.
(2004b)
in
coas al
dunes
o
Is ael.
I
was
simila
o
alues
egis-
e ed
by
Kollmann
e
al.
(2009)
o
he
alien
Rosa
ugosa
(16.4%;
18
yea s
analyzed)
in
coas al
dunes
o
Denma k.
The
annual
a e
o
la e al
expansion
o
he
R.
monospe ma
popula ion
(65.5
m
y −1)
was
also
significan ly
highe
han
he
la e al
expansion
a e
o
o he
woody
species
in
coas al
dunes
o
England
epo ed
by
Fulle
and
Boo man
(1977;
5
m
y −1 o
he
sh ubs
Rhododen-
d on
pon icum
and
Hippophäe
hamnoides).
The
obse ed
end
o
expansion
o
R.
monospe ma
in
coas al
dunes
o
sou he n
Spain
is
he e o e
compa able
o
he
in asi e- ype
expansion
exhibi ed
by
alloch honous
in asi e
species
(see
also
Buell
e
al.,
1995).
The
linea
end
o
inc ease
in
popula ion
densi y
shown
in
he
cen-
al
sec o
o
he
spi
could
be
ela ed
o
he
simul aneous
g ow h
o
a
majo i y
o
indi iduals
o
he
same
age,
delibe a ely
in o-
duced
o
fix
dunes
nea
o
he
unny
fishe y
ha
was
ac i e
in
his
sec o
du ing
he
fi s
hal
o
20 h
cen u y.
This
ac
mus
be
aken
in o
accoun
when
in e p e ing
he
esul s,
as
he
his-
o ical
mass
plan a ion
o
R.
monospe ma
ha
occu ed
he e,
means
ha
he
na u al
slow
in oduc ion
and
es ablishmen
phases
o
in asion
dynamics
did
no
occu
in
his
case.
Judging
abou
he
in asi e
po en ial
o
R.
monospe ma
i
mus
be
aken
in o
accoun
gene ally,
ha
his
species
is
also
used
as
an
o namen-
al
plan ,
a
ac
which
acili a es
i s
in oduc ion
in o
new
a eas
(Tala e a,
1999).
Resul ing
om
such
in oduc ions,
R.
monospe ma
is
p esen ly
conside ed
o
be
a
non-na i e
in asi e
plan
o
coas al
sh ublands,
g asslands,
and
he
chapa al
in
Cali o nia,
USA
(Cal-
IPC,
2006;
Randall,
1997;
Rejmanek
and
Randall,
1994),
and
an
in asi e,
noxious
weed
on
he
coas s
o
SW
Aus alia
(Randall,
2007).
The
eco ded
annual
g ow h
a e
o
R.
monospe ma
co e age
on
he
in es iga ed
spi
was
ound
o
be
9.8
imes
ha
which
Ku iel
e
al.
(2004a)
ound
o
he
o al
ege a ion
co e age
(1.8%;
34
y .)
in
plan -colonized
coas al
dunes
unde
Medi e anean
cli-
ma e
o
sou he n
Is ael.
In
ha
s udy,
he
expansion
o
ege a ion
was
ound
o
be
ela ed
o
he
dune
s abiliza ion
p ocess
and
o
a
dec ease
in
he
abundance
o
sand-dwelling
o ganisms.
The
expansion
o
R.
monospe ma,
and
in
consequence
he
p og essi e
s abiliza ion
o
dunes,
in
he
El
Rompido
spi
and
o he
dune
sys-
ems
in
SW
Spain,
ha e
nega i e
consequences
o
he
conse a ion
o
semi-s abilized
dunes
(Eu opean
Communi ies
p o ec ed
habi-
a
2130
and
2250;
Anonymous,
1992).
In
addi ion
o
he
apid
expansion,
he
species
could
induce
significan
changes
in
he
local
en i onmen
ha
may
acili a e
he
ec ui men
o
species
less
ole an
o
en i onmen al
s ess
(mainly
ni ophilous,
ude al
and/o
weed
species,
such
as
Anagallis
a ensis,
E ige on
suma en-
sis,
Pa onychia
a gen ea,
Solanum
ala um
and
U ica
memb anacea,
among
o he s).
Fu he ,
i
may
nega i ely
a ec
he
dune
flo a
benea h
he
canopy,
inc easingly
wi h
highe
R.
monospe ma
age
and
co e age
(Mu˜
noz
Vallés
e
al.,
2011;
Zunzunegui
e
al.,
2012).
This
significan
al e s
he
ege a ion
abundance,
composi ion
and
s uc u e,
wi h
nega i e
e ec s
upon
o he
dune
plan
species
he
i ali y
o
which
will
su e
om
he
decline
o
open,
semi-
s abilized
dunes
(Mu˜
noz
Vallés
e
al.,
2009,
2011).
In
his
way,
R.
monospe ma
beha es
as
a
na i e-in asi e
species,
and
shows
a
S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267 265
high
po en ial
o
inducing
significan
unc ional
changes
wi hin
he
dune
ecosys em.
In
ag eemen
wi h
Valé y
e
al.
(2009),
his
s udy
suppo s
he
idea
ha
he
expansion
o
na i e-in asi e
species
may
ha e
neg-
a i e
e ec s
on
ecosys em
di e si y
and
s abili y.
In
addi ion,
i
may
ha e
g ea
ele ance
o
he
conse a ion
o
p ope ies
and
p ocesses
wi hin
he
ecosys ems
unde
R.
monospe ma
in asion,
likewise
as
i
would
be
he
case
o
an
in asi e
sp ead
o
alien
species.
Excessi e
co e age
o
R.
monospe ma
should
he e o e
be
con-
olled
and
managed
in
some
loca ions
o
SW
Spain
and
S
Po ugal,
in
o de
o
p ese e
he
exis ing
plan
communi ies
and
he
pe -
sis ence
and
unc ioning
o
semi-s abilized
dune
habi a s.
In
his
ega d,
i
has
been
obse ed
ha
seawa e
inunda ion
by
high
ides
causes
ob ious
physiological
damage
o
his
na i e-in asi e
species,
and
sand
bu ial
and
sal wa e
in usion
appea
o
limi
i s
es ablishmen
(Mu˜
noz
Vallés
e
al.,
in
p ess).
I
has
been
also
obse ed
ha
R.
monospe ma
seedlings
and
young
s ands
de elop
success ully
in
open
a eas,
bu
ec ui men
unde
adul
canopies
o
he
species
is
e y
limi ed.
Rey
Benayas
e
al.
(2002)
ound
significan
compe i ion
be ween
weeds
and
R.
sphae oca pa
seedlings
when
g owing
oge he ,
bu
shading
(e.g.
p o ided
by
Re ama
adul s)
imp o ed
pe o mance
o
he
seedlings.
These
la -
e
seemed
o
be
in
a
compe i ion
o
wa e
esou ces
wi h
weeds.
This
same
in e ac ion
could
occu
be ween
Re ama
adul s
and
seedlings,
so
adul s
could
con ol
he
popula ion
ec ui men
unde
i s
canopies
h ough
compe i ion
–
bu
his
should
no
sol e
he
expansion
p oblem.
In
his
ega d,
u he
s udies
a e
equi ed
in
o de
o
assess
he
ac o s
ha
limi
seedling
es ablishmen
and
su i al.
P e ious
s udies
ha e
iden ified
wild
abbi s
as
impo an
dis-
pe se s
o
R.
monospe ma
in
coas al
dunes
(Dellafio e
e
al.,
2006,
2010).
I
is
es ima ed
ha
each
abbi
could
dispe se
be ween
2484
and
4041
seeds
y −1,
a
bo h
sho
and
long
dis ances
(up
o
335
m
om
he
nea es
ma u e
plan
o
R.
monospe ma),
con-
ibu ing
o
he
de elopmen
o
he
species
seed
bank.
In
addi ion,
passage
h ough
he
abbi
gu
imp o es
seed
ge mina ion
a es,
om
13%
o
24%,
(Dellafio e
e
al.,
2006).
These
p ocesses
ce -
ainly
ha e
acili a ed
–
and
con inue
doing
so
–
he
apid
la e al
expansion
o
he
R.
monospe ma
popula ion
o e
he
con inuous
dune
su ace.
They
con ibu e
also
o
o e come
obs acles
in
he
dune
su ace
ep esen ed
by
sal
ma shes
and
idal
swales
(Gallego-
Fe nández
e
al.,
2006).
As
in
p e ious
s udies
ound,
he
in e ac ion
be ween
wild
abbi s
and
R.
monospe ma
wo ks
as
a
eed-back
sys em,
whe e
he
abbi
ac s
as
seed
dispe se
o e
sho
and
long
dis ances
and
imp o es
seed
ge mina ion
a es
(Dellafio e
e
al.,
2006),
while
R.
monospe ma
in
u n
p o ides
abbi s
wi h
e uge
space
and
ood
(Dellafio e
e
al.,
2008).
Ou
esul s
show
a
long- e m
ne
inc ease
o
he
ex en
and
co e age
o
he
sh ub
popula ion
in
a eas
whe e
abbi s
a e
dis ibu ed,
wi hou
he
concu ence
o
ano he
ype
o
dis u bance,
a
si ua ion
which
leads
o
he
s abiliza ion
o
he
dune
sys em.
This
clea ly
con-
as s
wi h
p e ious
s udies
o
g azing
in
Eu opean
coas al
dunes,
whe e
abbi s
play
an
impo an
ole
in
main aining
low
le els
o
ege a ion
co e age
and
inc easing
sand
dune
mobili y
(Bu gg aa -
an
Nie op
and
an
de
Meijden,
1984;
Hodgkin,
1984;
Ranwell,
1960).
In
con as ,
sec o s
subjec ed
o
g azing
by
li es ock
showed
significan ly
lowe
inc eases
in
R.
monospe ma
co e age
o e
ime,
which
was
e en
lowe
in
he
a ea
subjec ed
o
highe
p essu e
(S1).
This
led
o
main enance
o
an
open
ype
o
plan
communi y
o
a
longe
pe iod
o
ime.
This
e a ding
e ec
on
expansion
and
den-
sifica ion
o
he
R.
monospe ma
popula ion
p obably
esul s
om
high
consump ion
a es
o
he
Re ama
ui s
by
goa s
and
sheep
(Zunzunegui
e
al.,
2012),
as
well
as
by
a
poo
capaci y
o
hese
animals
o
dispe sal.
Cas o
e
al.
(2005)
ound
ha
passage
o
Re ama
sphae oca pa
seeds
h ough
he
gu
o
li es ock
did
no
sig-
nifican ly
inc ease
ge mina ion
a es,
excep
whe e
he
seed
had
been
damaged
du ing
mas ica ion.
Howe e ,
we
ound
ha
pas-
sage
h ough
goa
diges i e
sys ems
(whe e
he
seed
emained
inside
he
gu
om
1
o
5
days),
inc eased
subsequen
ge mina-
ion
a es
om
5%
(in
non-consumed
seeds)
o
48%
(Mu˜
noz
Vallés,
unpublished
da a).
O e
a
fi e
days
obse a ion
pe iod
we
ound
a
final
32%
seed
eco e y
om
exc emen s
o
he
expe imen ally
s udied
goa s.
This,
oge he
wi h
he
final
ge mina ion
a e
da a
(weigh ed
o
mean
daily
eco e y),
esul ed
in
a
final
e ec i e
ge mina ion
pe cen age
o
16%,
which
was
simila
o
he
a es
in
con ol
seeds
(13%)
epo ed
by
Dellafio e
e
al.
(2006).
In
his
com-
pa ison,
i
is
impo an
o
ake
in o
accoun
he
loss
o
seeds
due
o
consump ion
and
subsequen
de eca ion
by
goa s
ou side
he
dune
sys em.
This
does
no
occu
o
he
same
ex en
in
he
case
o
seed
consump ion
by
abbi s,
as
hey
a e
esiden
wi hin
he
dune
sys em.
In
addi ion,
p eda ion
on
seedlings
o
R.
monospe ma
by
domes ic
li es ock
has
been
no
quan ified.
Ou
esul s
concu
wi h
hose
o
o he
s udies
whe e
li es ock
g azing
con ibu es
o
he
main enance
o
open- ype
plan
communi ies
as
well
as
o
he
mobili y
o
sands,
due
o
he
dec ease
in
ege a ion
co -
e age
(Kooijman
and
Van
de
Meulen,
1996;
Le in
and
Ben-Do ,
2004;
Manzano
and
Ná a ,
2000;
Zhao
e
al.,
2005).
In
his
way,
he
obse ed
co e age
inc ease
in
sec o
S2
om
1989
o
2001
could
be
ela ed
o
he
educ ion
in
in ensi y
o
pas u e
ac i i ies
ha
ollowed
he
decla a ion
o
he
si e
as
a
p o ec ed
a ea
(1989).
Acco ding
o
he
esul s
epo ed
by
o he
au ho s,
such
e ec s
o
li es ock
ac i i y
on
dune
ege a ion
can
change,
depending
on
ac o s
such
as
soil
cha ac e is ics
(Kohyani
e
al.,
2008;
Van
Dijk,
1992;
Zuo
e
al.,
2008),
in ensi y
o
g azing
(Ca e ,
1988),
o
also
he
moni o ing
scale
(Kohyani
e
al.,
2008).
In
some
cases,
dec eas-
ing
compe i i e
in e ac ions
caused
by
high
g azing
p essu e
ha e
esul ed
in
he
loss
o
g asslands
and
he
u he
de elopmen
o
sh ub
o ma ions
(Ise mann
e
al.,
2007;
Ro h
e
al.,
2009).
How-
e e ,
o he
nega i e
e ec s
o
li es ock
pas u e
ac i i ies
he e,
like
ampling,
soil
compac ion
and
eu ophica ion
o
dispe sion
o
ad en i ious
species
om
ou side
he
sys em
ha e
no
ye
been
quan ified.
Low
o
medium
co e age
o
R.
monospe ma
would
be
posi i e
o
he
su i al
o
he
endange ed
ep ile
Chamaeleo
chamaeleon,
ha
is
p esen
on
he
s udy
a ea,
bu
in ensi e
g az-
ing
and
ampling
by
domes ic
li es ock
would
be
e y
ha m ul
o
he
ep oduc ion
o
his
species,
since
hey
lay
hei
eggs
on
he
soil.
Implemen a ion
o
g azing
by
li es ock
may
appea
o
be
a
sui -
able
managemen
ool
o
he
con ol
o
excessi e
co e age
o
R.
monospe ma
in
hese
coas al
dunes,
and
i
has
been
ad oca ed,
indeed,
by
Plassmann
e
al.
(2010).
Howe e ,
he
po en ial
conse-
quences
o
adop ing
such
a
managemen
s a egy
in
a
agile
dune
ecosys em
need
o
be
unde s ood
mo e
ully
(Ra aele
e
al.,
2007).
Zunzunegui
e
al.
(2012)
in
a
s udy
a
he
El
Rompido
spi
in es-
iga ing
he
e ec s
o
di e en
he bi o y
in ensi y
and
densi y
o
R.
monospe ma
on
an
endange ed
sh ub
species,
Thymus
ca nosus,
discou age
he
managemen
wi h
li es ock.
They
an icipa e
he
u u e
ex inc ion
o
T.
ca nosus
in
he
dunes
because
i
su e s
om
he
compe i ion
wi h
R.
monospe ma,
al hough
he
speed
o
his
p ocess
would
be
condi ioned
by
he
li es ock
p essu e
in ensi y.
This
la e
s udy
p oposes,
in
e ms
o
T.
ca nosus
popula ion
con-
se a ion,
a
clea -cu ing
managemen
o
R.
monospe ma
plan s,
educing
by
his
way
hei
numbe
in
o de
o
main ain
open
dunes
spaces.
Acknowledgemen s
We
hank
he
Spanish
Minis y
o
En i onmen
(Coas al
Del-
ega ion)
o
unding,
and
he
s a
o
he
Ma ismas
del
Pied as
y
Flecha
Li o al
de
El
Rompido
p o ec ed
a ea
o
hei
help
and
266 S.
Mu˜
noz-Vallés
e
al.
/
Flo a
208 (2013) 259–
267
suppo .
Thanks
also
o
K.
MacMillan
o
ansla ion
and
e ision
o
he
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e sion
o
he
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