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HUMAN IMPACT ON PEATLANDS: FROM BIOGEOCHEMICAL ISSUES TOWARDS
SUSTAINABLE LAND USE OPTIONSAuthor | J.M.H. van Diggelen
HUMA
N IMP
ACT O
N PEAT
LANDS
: FR
OM BI
OGEO
CHEM
ICAL IS
SUES
TOWA
RDS
SUSTA
INABL
E LAN
D USE
OPTIO
NSAu
thor
| J.
M.H
. van
Dig
gele
n
Prin
ted
by: I
pska
mp
Druk
kers
, Ens
ched
e
ISBN
978
-94-
6259
-818
-8©
201
5 Jo
seph
a M
.H. v
an D
igge
len
The
rese
arch
pre
sent
ed in
this
thes
is w
as fu
nded
by
Radb
oud
Uni
vers
ity N
ijmeg
en a
nd B
-WAR
E Re
sear
ch C
entr
e.
HUMA
N IMP
ACT O
N PEAT
LANDS
: FR
OM BI
OGEO
CHEM
ICAL IS
SUES
TOWA
RDS
SUSTA
INABL
E LAN
D USE
OPTIO
NS
ter v
erkr
ijgin
g va
n de
gra
ad v
an d
octo
r aa
n de
Rad
boud
Uni
vers
iteit
Nijm
egen
volg
ens b
eslu
it va
n he
t col
lege
van
dec
anen
in
het
ope
nbaa
r te
verd
edig
en o
p w
oens
dag
4 no
vem
ber 2
015
om 1
2.30
uur
pre
cies
door
Jose
pha
Mar
ia H
endr
ika
van
Digg
elen
gebo
ren
op 1
5 m
ei 1
982
te S
chai
jk
Prom
otor
en:
Prof
. dr.
L.P.
M L
amer
sPr
of. d
r. J.G
.M. R
oelo
fs
Copr
omot
or:
Prof
. dr.
A.J.P
. Sm
olde
rs
Man
uscr
iptc
omm
issi
e:Pr
of. d
r. H.
de
Kroo
n Pr
of. d
r. A.
P. G
root
jans
Prof
. dr.
J.T.A
. Ver
hoev
en
Para
nim
fen:
M. P
oele
nG.
van
Dijk
in a
n ev
er c
hang
ing,
turb
ulen
t and
incr
easin
gly
crow
ded
wor
ld.
to d
evel
op so
me
good
stra
tegi
es to
pre
serv
e pe
atla
nds i
n th
e lo
ng te
rm.
Isn’t
it funny
how da
y by da
y noth
ing cha
nges, b
ut when
you loo
k back
everyth
ing is
differe
nt...
C.S.
Lew
is
Co
nten
ts
6 Ho
w to
cho
ose
betw
een
evils
: The
bio
geoc
hem
istry
of d
rain
ed c
oast
al
pe
atla
nds c
ause
s a m
anag
emen
t dile
mm
a
81
7 Ge
nera
l disc
ussio
n &
synt
hesis
9
3 8
Refe
renc
es
1
05 11
Da
nkw
oord
13
7
Jo
seph
a M
.H. v
an D
igge
len
Pe
atla
nds
they
also
incl
ude
tree
- or s
hrub
-dom
inat
ed s
wam
ps a
nd g
rass
-dom
inat
ed m
arsh
es (e
.g. M
itsch
&
Cype
race
aem
acro
phyt
es a
nd b
row
nmos
ses,
whi
le b
ogs
are
char
acte
rised
by
Spha
gnum
spe
cies
(Whe
eler
&
Proc
tor,
2000
; Lam
ers
et a
l
(O2
et a
l
Spha
gnum
poo
r et
al.,
2 in 1
80 c
ount
ries
(3%
Lim
pens
et a
lve
ry im
port
ant f
or h
uman
wel
fare
and
pro
vide
a h
igh
varie
ty o
f nat
ural
reso
urce
s and
serv
ices
, suc
h
et a
l., 1
997;
Mits
ch &
Gos
selin
k, 2
000;
Joos
ten
& C
lark
e, 2
002;
area
s to
impr
ove
agric
ultu
ral r
esou
rce
man
agem
ent.
This
serio
usly
thre
aten
s th
e cu
rren
t sta
tus
of
11
1
glob
al C
sto
rage
(e.g
. Cos
tanz
a et
al.,
199
7; M
itsch
& G
osse
link,
200
0; B
rinso
n &
Mal
váre
z, 20
02;
Zedl
er &
Ker
cher
, 20
05;
Bobb
ink
et a
let
al.,
200
3;
Bella
my
et a
lEu
rope
, Nor
th A
mer
ica, a
nd E
ast A
sia e
ven
mor
e th
an 8
0%, o
f the
wet
land
s hav
e be
en lo
st o
r sev
erel
y de
grad
ed (e
.g. J
oost
en &
Cla
rke,
200
2; L
aiho
, 200
6; L
amer
s et
al
th c
entu
ry,
et a
l.,
1213
Pe
atla
nds i
n th
e N
ethe
rland
sDu
ring
the
Holo
cene
, the
wes
tern
par
t of T
he N
ethe
rland
s bec
ame
cove
red
by m
arsh
land
as a
resu
lt of
incr
ease
d ra
infa
ll co
mbi
ned
with
poo
r dr
aina
ge d
ue to
the
mar
ine
clay
sub
soil.
Sev
eral
met
res
man
aged
to tr
ansf
orm
thes
e m
arsh
land
s in
to a
rabl
e la
nd b
y di
ggin
g ca
nals
and
ditc
hes
to e
nabl
e th
cen
tury
onw
ards
, pea
tland
coas
tal a
reas
of t
he N
ethe
rland
s, w
hile
fres
hwat
er p
eatla
nds l
ess r
ich
in S
can
be fo
und
mor
e in
land
. Cu
rren
tly, D
utch
pea
tland
s sho
w a
typi
cal p
olde
r lan
dsca
pe, i
nclu
ding
a la
rge
netw
ork
of w
ater
way
s su
rrou
ndin
g la
rge
drai
ned
peat
land
are
as.
Thes
e ar
eas
are
pred
omin
antly
use
d fo
r ag
ricul
tura
l
the
Dutc
h m
anag
ed p
eat m
eado
ws a
s bre
edin
g ar
eas o
n a
larg
e sc
ale,
and
are
now
aday
s pro
tect
ed
et a
l
et a
lbo
tani
cal a
nd zo
olog
ical
div
ersit
y th
an th
e ag
ricul
tura
l are
as. N
ever
thel
ess,
bot
h th
e se
mi-n
atur
al
the
cultu
ral h
erita
ge o
f a la
rge
part
of t
he N
ethe
rland
s. In
tens
ive
drai
nage
has
larg
ely
decl
ined
the
rate
s up
to 1
cm
per
yea
r in
the
rem
aini
ng a
reas
(Oud
e Es
sink
et a
l., 2
010;
Hoo
glan
d et
al
high
er th
an th
e in
tens
ivel
y dr
aine
d po
lder
s or u
rban
are
as su
rrou
ndin
g, w
hich
has
stro
ngly
alte
red
et a
l., 1
996;
Koe
rsel
man
& V
erho
even
, 19
92; B
eltm
an e
t al.,
200
1; L
amer
s et a
l
Bi
ogeo
chem
ical
issu
es in
pea
tland
s
Alte
red
hydr
olog
y
low
ered
wat
er le
vels
and
wat
er s
hort
age
(e.g
. Mid
dlet
on e
t al.,
200
6a; L
amer
s et
al.,
200
2b; 2
014;
tem
pera
ture
s will
incr
ease
with
1 to
5C
in th
e 21
st
2009
; IPC
C, 2
014;
Hay
et a
l
1
2 em
issio
ns t
o th
e at
mos
pher
e an
d co
ncom
itant
la
nd s
ubsid
ence
(e.g
. Kas
imir-
Klem
edts
son
et a
l., 1
997;
Lai
ho, 2
006;
Lam
ers
et a
let
al
2 in t
he p
eat
is an
impo
rtan
t
& P
roct
or, 2
000;
Lam
ers e
t al
or s
emi-t
erre
stria
l pea
tland
s (H
ogg
et a
l., 1
995;
Koo
ijman
& P
aulis
sen,
200
6; K
limko
wsk
a et
al.,
20
07; G
eurt
s et
al.,
200
9; L
amer
s et
al.,
Roel
ofs,
1995
; Lam
ers e
t al
et a
l
1996
; Old
e Ve
nter
ink
et a
ldo
min
ance
of g
ram
inoi
ds, w
hich
cau
ses b
iodi
vers
ity lo
ss in
wet
land
s (Bo
bbin
k et
al
will
dire
ctly
incr
ease
nitr
ate
(NO
3-43-
14
Dutc
h co
asta
l pea
tland
are
a, lo
cate
d se
vera
l met
ers b
elow
sea
wat
er le
vel
34+
2
32SO
4et
al.,
42-
4+ and
NO
3-
deriv
ed fr
om th
e sy
stem
itse
lf (S
mol
ders
et a
l42-
P re
leas
e bo
th fr
om a
naer
obic
pea
t so
ils o
r un
derw
ater
sed
imen
ts, a
s m
icro
bial
SO
42-
rate
s can
be
enha
nced
. As a
resu
lt of
SO
42-2-
2+ a
nd m
obili
ses
P (L
amer
s et
al.,
200
2a, S
mol
ders
et a
l42-
in t
he p
eat
soil,
or
by t
he s
uppl
y of
SO
42-
2
such
as
NO
3- or S
O42-
and
P bo
und
in th
e or
gani
c so
il. In
are
as w
ith se
epag
e, S
O42-
and
NO
3- leve
ls m
ay a
lso in
crea
se a
s a
resu
lt of
the
disc
harg
e of
gro
undw
ater
from
surr
ound
ing
fore
sts o
r agr
icul
tura
l are
as.
et a
l., 1
998b
; Sm
olde
rs et
al
et a
l3-
2+2+
rich
wat
er in
fens
dec
reas
es a
s th
e th
ickn
ess
of t
he p
eat
mat
incr
ease
s, w
hich
lead
s to
nat
ural
et a
l.,
Spha
gnum
. Thi
s ge
nera
tes
a lo
ss o
f
15
1
typi
cal f
en s
pecie
s an
d a
decr
ease
of o
vera
ll bi
odiv
ersit
y in
fens
(Bel
tman
et a
l., 2
001;
Mid
dlet
on e
t al
., 20
06a;
Lam
ers e
t al
To
xici
tySe
vera
l com
poun
ds th
at a
re u
sual
ly fo
und
in w
ater
logg
ed, a
naer
obic
pea
t soi
ls su
ch a
s NH 4+ , S
2- or
2+
et a
l., 1
995;
Luc
asse
n et
al
., 20
02; L
amer
s et
al
4+ is th
e do
min
ant N
form
an
d is
easil
y ta
ken
up a
s a n
utrie
nt. A
t hig
h le
vels,
how
ever
, NH 4+
et a
l
2 rel
ease
fro
m t
he r
oots
, the
reby
enh
anci
ng o
vera
ll et
al
stro
ng la
nd s
ubsid
ence
in th
ese
coas
tal a
reas
may
also
enh
ance
sal
ine
seep
age
wat
er to
intr
ude
wat
er s
yste
ms
(Zed
ler &
Ker
cher
, 200
5; O
ude
Essin
k et
al.
(Nie
lsen
et a
l., 2
003;
Her
bert
et a
l-
(SO
42-2+
et a
l
et a
l., 2
003;
Wes
ton
et a
let
al
Di
lem
mas
for p
eatla
nd m
anag
emen
t
16
et a
ldi
lem
mas
. Wat
er le
vels
shou
ld b
e hi
gh to
pre
vent
land
sub
siden
ce, w
hile
farm
ers
pref
er a
low
er
Alth
ough
ther
e is
ampl
e kn
owle
dge
abou
t the
mai
n pr
oble
ms a
nd th
reat
s in
peat
land
s, th
e un
derly
ing
The
rese
arch
incl
uded
in th
is th
esis
was
real
ised
at B
-WAR
E (B
ioge
oche
mic
al W
ater
Man
agem
ent
the
Radb
oud
Uni
vers
ity N
ijmeg
en. A
t B-W
ARE
Rese
arch
Cen
tre,
app
lied
rese
arch
is c
arrie
d ou
t to
thos
e in
The
Net
herla
nds.
ch
ange
s. A
nov
el c
ontr
ollin
g m
echa
nism
was
sug
gest
ed, w
hich
may
hel
p to
dev
elop
mea
sure
s to
4+
( a
nd
resu
lts m
ay c
ontr
ibut
e to
impr
ove
futu
re w
etla
nd m
anag
emen
t in
orde
r to
cons
erve
bio
dive
rsity
.
17
1
disc
usse
d. C
hapt
er 7
pro
vide
s a
synt
hesis
whi
ch c
ombi
nes
our
resu
lts a
s de
scrib
ed in
thi
s th
esis
natu
ral a
nd a
gric
ultu
ral p
eatla
nds i
n th
e lo
ng te
rm.
1819
1
2 |
New
insi
ghts
into
pho
spho
rus
Ab
stra
ct
betw
een
salin
ity c
hang
es a
nd n
utrie
nt c
yclin
g in
fres
hwat
er s
yste
ms
are
not f
ully
und
erst
ood.
To
2+
2+ i
mm
obili
ses
P, e
ven
thou
gh i
t is
2+
2+
-ric
h se
dim
ents
, its
2+
and
O2.
21
2
mon
opol
ising
ligh
t (Sm
ith, 2
003;
Lam
ers e
t al.,
200
2a; G
eurt
s et a
l
(Nie
lsen
et a
let
al.,
200
0; Jo
rdan
et
al
loss
of b
iodi
vers
ity.
et a
l., 1
988;
Car
aco
et a
l., 1
989;
Lam
ers e
t al.,
199
8a;
Smol
ders
et
al.
(O2
2+
the
surfa
ce w
ater
.
42-
Cara
co e
t al.,
199
3; S
mol
ders
and
Roe
lofs
, 199
5; S
mol
ders
et a
l., 2
006a
; Hup
fer a
nd L
ewan
dow
ski,
2008
; Geu
rts
et a
l2-
2+ in
sed
imen
t por
e w
ater
, 2+
2+
et al
et al
.
-1
242-
enr
iched
w
etla
nds (
Cara
co e
t al.,
198
9; Sm
olde
rs e
t al.,
200
1; Sm
olde
rs e
t al.,
200
6a; H
upfe
r and
Lew
ando
wsk
i,
42-
et a
l., 2
002a
; Sm
olde
rs e
t al.,
200
6a; G
eurt
s et a
l
42-42-
avai
labi
lity
Cl- a
nd S
O42-
bind
ing
sites
(Car
aco
et a
l., 1
989;
Bel
tman
et a
lal
so le
ads t
o in
crea
sed
Ca2+
2+ a
nd ca
lciu
m ca
rbon
ate
(CaC
O3
et a
l., 1
988;
Hou
se e
t al.,
19
99; N
ielse
n et
al
et a
l., 2
001;
Geu
rts e
t al.,
201
0;
22
Wu
et a
l
-ric
h se
dim
ents
from
a c
oast
al
fres
hwat
er w
etla
nd w
ere
subj
ecte
d to
thr
ee n
atur
ally
occ
urrin
g w
ater
typ
es c
hara
cter
ised
by
(Lam
ers e
t al
et a
l.,
1989
; Sm
olde
rs e
t al.,
200
1, S
mol
ders
et a
l., 2
006a
; Hup
fer a
nd L
ewan
dow
ski,
2008
; Geu
rts e
t al.,
of se
dim
ents
com
mon
for c
oast
al w
etla
nds.
M
ater
ial a
nd M
etho
ds
Sam
plin
g ar
eaIn
this
stud
y, pe
at se
dim
ents
wer
e us
ed fr
om th
e co
asta
l low
land
fen
area
Wor
mer
- and
Jisp
erve
ld
syst
em. T
he p
eatla
nd c
ompr
ises
ca. 5
00 h
a of
ope
n w
ater
and
ca.
166
0 ha
of
peat
mea
dow
s,
Ex
perim
enta
l des
ign
On
18 M
arch
200
8, tw
o ty
pes o
f sub
mer
ged
peat
sedi
men
t wer
e co
llect
ed fr
om a
ditc
h at
a d
epth
23
2
To m
inim
ise O
2 intr
usio
n, th
e se
dim
ents
wer
e st
ored
ana
erob
ical
ly a
t 4o C
in la
rge,
clos
ed co
ntai
ners
.
envi
ronm
enta
lly re
leva
nt te
mpe
ratu
re o
f 15o
open
cyl
inde
r sys
tem
was
use
d.
trea
tmen
ts: r
ainw
ater
(low
sal
inity
; 100
μm
ol C
l L-1
-1
and
fres
hwat
er (c
ontr
ol; 7
mm
ol C
l L-1
et a
l
star
ted
by r
epla
cing
the
con
trol
wat
er w
ith t
he a
ppro
pria
te t
reat
men
t w
ater
dur
ing
4 w
eeks
, to
wat
er d
urin
g th
is st
agna
nt p
erio
d.
24
2
2 to th
e su
rface
wat
er. D
urin
g bo
th
Ch
emic
al a
naly
ses
cont
aini
ng 0
.2 M
H3PO
42
2di
rect
ly tr
ansp
orts
the
CO2
3-3-
2+ ra
ther
than
far l
ess
3+. T
he r
emai
ning
sam
ples
wer
e st
ored
at
-20°
C in
ord
er t
o de
term
ine
the
follo
win
g io
n 3- (K
amph
ake
et
al4+
43--
Na+ a
nd K
+
200
mg
dry
sedi
men
t w
as d
iges
ted
in 5
mL
HNO
32O
2
25
2
Et
hics
stat
emen
t
faci
litat
ed t
his
prog
ram
me
and
the
natu
re m
anag
emen
t au
thor
ity ‘
Nat
uurm
onum
ente
n’ g
ave
perm
issio
n to
take
sam
ples
in th
eir r
eser
ve.
Re
sults
Po
re w
ater
che
mist
ry
+
wat
er N
a++
sedi
men
t typ
e w
as fo
und
for b
oth
Na+
wer
e fo
und
betw
een
sedi
men
ts o
r tre
atm
ents
(ave
rage
val
ues
rang
ed b
etw
een
0-50
μm
ol L
-1 fo
r se
dim
ent A
, and
bet
wee
n 0-
500 μm
ol L
-1
26
As a
resu
lt of
a h
ighe
r sal
inity
, Ca2+
incr
ease
d po
re w
ater
Ca2+
-1
high
sal
inity
tre
atm
ent
and
Ca2+
2+
2+
2+
betw
een
trea
tmen
ts w
ere
foun
d fo
r sed
imen
t B. P
ore
wat
er H
CO3-
of P
in p
ore
wat
er w
as fo
und
for
the
cont
rol t
reat
men
t, co
mpa
red
to t
he h
igh
and
low
sal
inity
27
2
Su
rfac
e w
ater
che
mist
ryA
high
er sa
linity
led
to g
radu
ally
incr
ease
d N
a+
+
2+
high
sal
inity
trea
tmen
t com
pare
d to
the
low
sal
inity
trea
tmen
t in
both
sed
imen
ts a
t a s
hort
term
28
term
. At s
edim
ent B
, how
ever
, no
chan
ge o
f P in
the
surfa
ce w
ater
was
foun
d fo
r any
of t
he sa
linity
trea
tmen
ts.
Ae
robi
c ve
rsus
ana
erob
ic su
rfac
e w
ater
The
O2
2
29
day
betw
een
trea
tmen
ts a
re
2
Di
scus
sion
-2 d
-1
et a
l15
0 μm
ol m
-2 d
-1
Geur
ts e
t al
or r
esus
pens
ion
(Suz
umur
a et
al
et a
l
et a
l2+
-1
(Car
aco
et a
l2-
et a
l., 2
002a
; Sm
olde
rs e
t al
This
imm
edia
te d
rop
of P
obs
erve
d up
on a
cha
nge
of th
e su
rface
wat
er c
hem
istry
str
ongl
y po
ints
2+ o
r Ca
CO3
et a
l., 1
988;
N
ielse
n et
al
2+
et a
l
et a
l2+
2an
d po
ssib
ly a
lso th
e pr
esen
ce o
f mic
robi
al m
ats (
Dupr
az e
t al
3 at t
he se
dim
ent s
urfa
ce, a
s HCO
3-
2-.
2+
et a
l-1
et a
l.,
2001
; Geu
rts e
t al
30
3+
et a
l., 1
980;
Hup
fer
et a
l., 1
988;
Hou
se e
t al.,
grow
ing
on to
p of
the
sed
imen
t (D
eine
ma
et a
l., 1
980;
Hup
fer
and
Uhlm
ann,
199
1; D
upra
z et
al.,
et a
l., 2
002;
et
al.,
200
4; G
eurt
s et a
l2+
at a
2
et a
l2
2 was
ava
ilabl
e in
the
surfa
ce
low
sal
inity
trea
tmen
t, w
here
no
S w
as a
dded
, ind
eed
show
ed th
at S
O42-
is b
eing
mob
ilise
d fr
om
2+
2 in
redu
ced
sedi
men
ts m
ay
2+
2
da
y
31
2
-rich
sed
imen
ts
2 ava
ilabi
lity
to p
reve
nt
et a
l
et a
l., 2
004;
Jord
an e
t al
42-
in b
oth
surfa
ce w
ater
and
por
e w
ater
and
a c
onsid
erab
le p
art
may
be
redu
ced
deep
er in
the
2-
will
reac
t with
O2
2+
2+ to
pre
vent
the
rele
ase
of P
to
the
surfa
ce w
ater
(Hup
fer a
nd L
ewan
dow
ski,
2008
; Geu
rts
et a
lan
d P
cycl
e (C
arac
o et
al
sedi
men
t int
erfa
ce. I
n se
dim
ent A
, O2
O2
2+
due
to th
e hi
gh C
l-et
al.,
198
9; B
eltm
an e
t al
2+
et a
lsu
rface
with
O2
2+
incr
ease
d SO
42-
2 42-
pla
y
32
0 C an
d
2 ra
tes an
d lo
wer
solu
bilit
y of
O2
will
lead
to st
rong
ly d
ecre
ased
O2
also
foun
d by
Sm
olde
rs e
t al.
or c
yano
bact
eria
may
dev
elop
, whi
ch w
ill fu
rthe
r de
crea
se t
he O
2 -ric
h se
dim
ents
that
show
ver
y 2+
4
33
2
Conc
lusi
ons
Alth
ough
thes
e Fe
S
in th
e sh
ort t
erm
,
, SO
4. I
n th
e lo
nger
term
,
3435
2
Pl
ant B
iolo
gy, a
ccep
ted
Ab
stra
ctEn
hanc
ed so
il am
mon
ium
(NH 4+
4+ is r
eadi
ly t
aken
up
as a
nut
rient
, sev
eral
4+
4+
wet
land
bio
dive
rsity
. Th
e re
spon
ses
of t
wo
wet
land
gra
min
oids
,an
d , t
o hi
gh (2
mm
ol L-1
-14+
4+
of N
take
n up
by
this
was
onl
y 10
% fo
r . A
lthou
gh b
oth
spec
ies
appe
ared
to b
e w
ell a
dapt
ed to
hig
h N
H 4+ load
ings
in th
e sh
ort t
erm
,
4+ load
ings
.
37
3
et a
l
the
mob
ile N
-form
nitr
ate
(NO
3-
et a
l.,et
al
Bobb
ink
et a
l., 1
998
et a
l. 20
01; G
eurt
s et
al.,
In w
etla
nd s
oils,
NO
3-
Inst
ead,
the
mor
e im
mob
ile N
H 4+
4+3-
diss
imila
tory
NO
3-4+
4+
Alth
ough
NH 4+ is
read
ily ta
ken
up as
a nu
trie
nt, v
ario
us sp
ecie
s fro
m d
ry, m
oist
and
wet
land
ecos
yste
ms
4+
et a
let
al
4+
et a
l., 2
008;
Van
der
Hei
de e
t al
et
al.,
2011
; Ste
vens
et a
let
al
4+
amon
g sp
ecie
s and
eve
n am
ong
plan
ts b
elon
ging
to th
e sa
me
spec
ies (
Cruz
et a
l.
4+ rich
, aci
dic
syst
ems
are
mor
e to
lera
nt to
hi
gh N
H 4+
4+ (De
Graa
f et a
let
al.,
et a
l4+
even
abs
ent
et a
l., 2
002;
Van
den
Ber
g et
al
wet
land
incr
ease
d N
H 4+
In th
e pr
esen
t stu
dy w
e th
eref
ore
test
ed th
e re
spon
ses
and
tole
ranc
es to
gre
atly
enh
ance
d N
H 4+ av
aila
bilit
y fo
r m
arsh
es,
and
Bro
wn
sedg
e (
test
ed w
heth
er th
ese
wet
land
spec
ies i
ndee
d pr
efer
the
upta
ke o
f NH 4+ ra
ther
than
NO
3- . Sec
ondl
y, th
eir
resp
onse
s an
d to
lera
nces
to
stro
ngly
inc
reas
ed N
H 4+
4+ tole
ranc
e of
wet
land
4+ lo
adin
gs to
, a
nd in
crea
sed
dom
inan
ce o
f J.
of sm
alle
r sed
ges s
uch
as
has
bee
n re
gula
rly o
bser
ved
in w
et
38
to h
ighl
y in
crea
sed
NH 4+
chan
ges a
nd to
mai
ntai
n w
etla
nd b
iodi
vers
ity.
M
ater
ial a
nd M
etho
ds
Expe
rimen
tal d
esig
n
and
Brow
n se
dge
- w
ere
sele
cted
. The
se sp
ecie
s usu
ally
occ
ur o
n m
oist
or w
et so
ils
pH o
f 5 -
6, a
nd c
an b
e ch
arac
teriz
ed a
s m
esot
roph
ic to
mod
erat
ely
eutr
ophi
c. In
the
Net
herla
nds,
NH 4+
-1
spec
ies (
Lam
ers e
t al
and
w
ere
colle
cted
from
a w
et
mea
dow
in a
nat
ure
rese
rve
near
Nijm
egen
, the
Net
herla
nds
(De
Bruu
k, 5
1o 46’ N
; 5o
into
an
aera
ted
18
3- or N
H 4+
adju
stm
ent o
f 7 d
ays
in a
erat
ed 1
8 bo
th N
O3- an
d N
H 4+10
0 μm
ol L
-1 C
a2+,
100 μm
ol L-1
Mg
2+, 2
00 μ
mol
L-1 K
+ , 100
μm
ol L-1
SO
42-, 2
00 μ
mol
L-1 P
O43-
, 0.8
μm
ol L-1
Mn2+
, 0.8
μm
ol
L-1 H
3BO3, 0
.7 μ
mol
L-1
Zn2+
, 0.2
7 μm
ol L
-1-1
Cu2+
, 0.0
08 μ
mol
L-1
M
o (D
e Gr
aaf e
t al.,
, with
bot
h N
O3- (2
0 μm
ol L
-14+ (2
0 μm
ol L
-1
,
and
1-cy
angu
anid
ine
( w
as a
dded
to th
e nu
trie
nt
(Va
n de
n Be
rg e
t al
.,cl
imat
e co
ntro
lled
room
with
a 1
6:8
hour
s da
y:ni
ght
regi
me,
at
an a
vera
ge te
mpe
ratu
re o
f 20°
C
phot
ons m
-2 s-1
. Thr
ee in
divi
dual
s of t
he sa
me
spec
ies w
ere
plac
ed in
eac
h pl
ant c
onta
iner
by
usin
g ei
ther
Na15
NO
3 or
15N
H 4Cl (
of to
tal N
2 ho
urs
of la
belin
g, p
lant
s w
ere
harv
este
dof
enr
iche
d N
in th
is pe
riod.
39
3
aer
ated
18
plan
ts o
f bot
h
Two
indi
vidu
als o
f the
sam
e sp
ecie
s wer
e pl
aced
in e
ach
In t
otal
, 32
plan
t co
ntai
ners
w
ere
used
. as
des
crib
ed a
bove
, and
last
ed f
or e
ight
wee
ks.
trea
tmen
ts: 2
0 μm
ol L
-1 N
H 4+ and
2 m
mol
L-1
NH 4+
usin
g 1.
2 m
ol L-1
HCl
and
1 m
ol L-1
The -1
1-c
yang
uani
dine
(4+
was
add
ed to
all
stor
age
tank
s (V
an d
en B
erg
et a
l.,
Ch
emic
al a
naly
sis
shoo
ts w
as d
irect
ly fr
ozen
at -
80o
2
Root
and
sho
ot m
ater
ial
was
drie
d fo
r 24
hou
rs a
t 70
o
(Mile
ston
e In
c.
15
15
amou
nt o
f 15N
.
40
Data
of s
ub-r
eplic
ates
from
the
2 or
3 p
lant
s gr
owin
g in
one
con
tain
er w
ere
pool
ed, l
eavi
ng 4
or
Re
sults
4+ ver
sus N
O3
Both
a
nd
wer
e ab
le to
take
up
NH 4+
as w
ell a
s N
O3- at
bot
h pH
4 a
nd p
H 6 4+
than
for N
O3-
4+3- in
a
nd
4+
spec
ies,
incr
ease
d N
H 4+
-l NH 4+
-1
NH4+
betw
een
pH a
nd h
igh
NH 4+ w
as fo
und
in s
hoot
bio
mas
s. Th
is le
d to
an
incr
ease
of s
hoot
bio
mas
s in
th
e hi
gh N
H 4+ tr
eatm
ent c
ompa
red
to th
e lo
w N
H 4+
4+
4+ and
pH
was
41
34
tota
l DW
h.
3
,
As a
resu
lt of
the
high
NH 4+
-l N
H 4+
at h
igh
NH 4+ ,
4+ tr
eatm
ent i
n sh
oot
t
reat
ed w
ith
high
NH 4+ . I
n ro
ot b
iom
ass
of 4+ a
nd h
igh
pH.
sho
wed
42
L4
4, a
t
shoo
t bio
mas
s, rh
izom
e bi
omas
s
43
mol L4mol L4
3
4+ and
pH,
and
an
incr
ease
in sh
oots
onl
y w
ith h
igh
NH 4+
, pH
and
NH 4+
4+ and
low
pH.
4+
4+ and
lo
w p
H.
4+
at l
ow p
H, re
gard
less
of t
he N
trea
tmen
t. M
agne
sium
4+ a
nd a
t a
4+ trea
tmen
ts w
ere
foun
d.
an in
crea
se o
f sho
ot b
iom
ass
at h
igh
NH 4+
et a
lbe
cam
e lim
ited
only
for
at b
oth
pH v
alue
s.
44
44
Both
spe
cies
res
pond
ed t
o en
hanc
ed N
H 4+
(R2
avai
labl
e N
. The
str
onge
st r
espo
nse
was
sho
wn
by, w
ith 7
3-75
% o
f th
e to
tal a
min
o 4+ tr
eatm
ent.
The
dom
inan
t N
-ric
h fr
ee a
min
o ac
id w
as a
spar
agin
e, w
hich
acc
ount
ed a
lread
y fo
r 71-
73%
of t
he to
tal f
ree
amin
o
. Alth
ough
show
ed a
sim
ilar
resp
onse
to
enha
nced
NH 4+
. J.
incr
ease
d le
ss th
an in
4+ an
d pH
. How
ever
,
4+ , whi
le th
is re
spon
se w
as a
bsen
t for
.
45
4
3
Di
scus
sion
The
upta
ke o
f NO
3- is a
n en
ergy
dem
andi
ng p
roce
ss fo
r pla
nts,
bec
ause
it h
as to
be
redu
ced
to N
H 4+ Be
caus
e N
H 4+
4+ ric
h en
viro
nmen
ts,
a pr
efer
ence
of
NH 4+
, and
to
a 4+ . A
lthou
gh N
H 4+ upt
ake
was
hig
her,
NO
3- w
as a
lso t
aken
up
read
ily, e
spec
ially
by
.4+ is
the
mos
t
et a
l.,et
al
plan
ts m
ight
faci
litat
e th
e up
take
of e
ither
NO
3- or N
H 4+ (Cha
ng e
t al
3- , also
has
the
high
est r
adia
l et
al
3-4+ a
re a
lso
4+ tre
atm
ent
(2 m
mol
L-1
Acor
us c
alam
uset
al.
and
(Tyl
ova
et a
l.,4+ lo
adin
gs
(res
p. 1
4.8
& 3
.7 m
mol
L-1
and
in
4+ . T
his a
bilit
y to
cha
nge
only
bio
mas
s 4+ t
oler
ance
. Suc
h a
tole
ranc
e to
hi
gh N
H 4+ load
ings
in w
etla
nd g
ram
inoi
ds w
as so
far o
nly
foun
d fo
r Phr
agm
ites a
ustr
alis
(Tyl
ová
et
al-1
we
foun
d fo
r bo
th s
peci
es, i
t see
ms
likel
y th
at g
row
th w
as N
-lim
ited
in th
e lo
w N
H 4+ trea
tmen
ts
(Was
sen
et a
l., 1
995;
Koe
rsel
man
& M
eule
man
, 199
6; O
lde
Vent
erin
k et
al.
high
NH 4+
4+ ava
ilabi
lity
incr
ease
d
How
ever
, hig
h N
H 4+
NH 4+ (O
lde
Vent
erin
k et
al.,
usin
g th
e m
etho
d of
Tim
mer
et
al
. The
se re
sults
are
in a
gree
men
t with
Tyl
ová
et a
lw
ho a
lso fo
und
indu
ced
K sh
orta
ge in
the
wet
land
spe
cies
a
s a
resu
lt of
hig
h N
H 4+ leve
ls.
yie
lded
a lo
wer
sho
ot b
iom
ass
4+
redu
ced
grow
th o
f (S
mol
ders
et a
l
4+
1987
; Luc
asse
n et
al.,
200
2; V
an d
en B
erg
et a
lup
take
of N
H 4+ , bec
ause
it is
acc
ompa
nied
by
the
rele
ase
of p
roto
ns t
o th
e rh
izosp
here
(Mill
er &
et
al
Cirs
ium
diss
ectu
m4+ a
t pH
4, b
ut n
ot a
t pH
6, d
ue to
inte
rnal
4+ u
ptak
e ra
tes
of b
oth
test
ed g
ram
inoi
ds w
ere
4+ is m
ost
46
incr
easin
g N
H 4+,
Callu
na v
ulga
ris,
(Van
den
Ber
g et
al.,
in R
icinu
s co
mm
unis
(Van
Beu
siche
m e
t al.,
(HCO
3-3-
are
unlik
ely
to h
ave
indu
ced
this
resp
onse
.
N
utrie
nt o
r tox
in?
Biom
ass
and
reso
urce
4+
per s
e in
ord
er to
incr
ease
thei
r
4+ . On
the
othe
r han
d, s
uch
chan
ges
in p
lant
mor
phol
ogy
and
phys
iolo
gy a
s a re
sult
of in
crea
sed
NH 4+
et a
l A
lthou
gh a
redu
ced
root
:sho
ot
NH 4+
et a
l.,,
foun
d fo
r bot
h gr
amin
oids
in re
spon
se to
enh
ance
d N
H 4+ up
take
(Van
Beu
siche
m e
t al.
& K
ronz
ucke
r, 20
02; T
ylov
a et
al.,
200
8; S
teve
ns e
t al Mar
schn
er, 1
995;
Sm
olde
rs e
t al
Kron
zuck
er, 2
002;
Mill
er &
Cra
mer
, 200
4; S
teve
ns e
t al.,
201
1fr
om N
ove
rload
. NH 4+
4+ dire
ctly
(Van
Dijk
& R
oelo
fs, 1
988;
Rab
e, 1
990;
Näs
holm
et a
l., 1
994;
Sm
olde
rs e
t al.,
200
0; T
omas
sen
et
alIn
our
stud
y,ac
ids
in a
bove
-gro
und
biom
ass
than
et
al
subj
ecte
d to
hig
h N
H 4+ leve
ls. In
our
stud
y, a
lso p
redo
min
antly
in
vest
ed in
the
amin
o ac
id a
spar
agin
e, w
hich
is k
now
n to
be
a st
orag
e co
mpo
und
and
the
maj
or
et a
l., 1
996;
Lea
et
al.
grow
ing
in a
N-r
ich
med
ium
, esp
ecia
lly w
hen
an o
ther
min
eral
ion,
pro
babl
y K
or C
a in
our
stu
dy,
beco
mes
lim
ited
for g
row
th (R
abe,
199
0; Le
a et
al
did
accu
mul
ate 47
3
4+ upt
ake,
whi
le
grow
th w
as n
ot e
nhan
ced,
how
ever
, ac
id N
com
poun
ds th
at w
ere
not m
easu
red
in o
ur st
udy.
Cru
z et
al.
plan
t tol
eran
ce to
NH 4+
et a
l.re
spon
ses
betw
een
wet
land
spe
cies
upo
n hi
gh N
H 4+
et a
lN
H 4+ to
lera
nce
of
, alth
ough
this
was
not
stud
ied
in th
is re
sear
ch. O
ur re
sults
impl
y th
at
4+
J. in
the
long
term
.
48
Conc
lusi
ons
4
4 lo
adin
gs, i
nclu
ding
4
4
49
3
in
wet
land
s: a
revi
ew
Leon
P.M
. Lam
ers,
Jose
pha
M.H
. van
Dig
gele
n, H
uub
J.M. O
p de
n Ca
mp,
Ab
stra
ct
biog
eoch
emica
l and
bio
logi
cal k
ey fa
ctor
s dr
ivin
g he
tero
gene
ity a
nd to
tal (
i.e.,
micr
oorg
anism
s an
d
com
bine
d ap
proa
ch h
as b
ecom
e ev
en m
ore
urge
nt.
51
4
et
al
-4 m
or
less
to 1
0-1 m
. One
sho
uld
real
ize th
at b
ecau
se th
e ro
ot sy
stem
s of
pla
nts
can
be v
ery
larg
e, u
p to
112
km
per
m2 o
f soi
l (Ro
bins
on e
t al
-3 m
and
101 m
(o
r ev
en d
eepe
r fo
r tr
ees;
Can
adel
l et a
l
an in
tegr
ated
app
roac
h of
mic
robi
olog
ists
and
plan
t ec
olog
ists,
in o
rder
to
unde
rsta
nd w
etla
nd
biog
eoch
emist
ry, b
iodi
vers
ity a
nd r
esili
ence
with
res
pect
to
ecos
yste
m c
hang
es. T
he p
urpo
se o
f
52
Unv
eget
ated
wet
land
soils
, suc
h as
dee
per p
arts
of l
ake
sedi
men
ts o
r sed
imen
ts o
f tur
bid
shal
low
chal
leng
es in
mic
robi
al e
colo
gy is
ther
efor
e to
incl
ude
this
hete
roge
neity
in re
sour
ces
(nut
rient
s,
et a
l., 2
000;
Bod
elie
r, 20
03; B
odel
ier
et a
let
al
As a
resu
lt of
het
erog
enei
ty, t
he a
vaila
bilit
y of
ele
ctro
n ac
cept
ors (
e.g.
, O2, N
O3-
3+, S
O42-
et a
l
N2
inco
mpa
rabl
e to
thos
e in
clo
sed
batc
h sy
stem
s.
et a
l
(Jack
son
& A
rmst
rong
, 199
9; V
isser
et a
l., 2
000a
,b; V
oese
nek
et a
l
2 in
the
uppe
r soi
l lay
er to
ens
ure
O2
et a
l., 1
989;
53
4
orga
nic
com
poun
ds a
nd m
iner
als,
car
ried
out
by fa
una
and
mic
roor
gani
sms,
str
ongl
y de
term
ine
that
sho
w a
hig
h av
aila
bilit
y of
N, e
.g.,
as a
resu
lt of
hig
h at
mos
pher
ic N
inpu
t. Ho
wev
er, s
tudi
es
show
muc
h in
cons
isten
cy (
Kalb
itz e
t al
et
al
et a
l
54
and
grow
th o
f bot
h pl
ants
and
mic
roor
gani
sms.
The
se e
lem
ents
are
pre
dom
inan
tly t
aken
up
by
plan
ts fr
om th
e so
il vi
a th
e ro
ots,
as o
nly
few
pla
nt sp
ecie
s may
take
up
high
am
ount
s of n
utrie
nts
1991
; Lar
cher
et a
lan
d bi
odiv
ersit
y. E
nhan
ced
N a
vaila
bilit
y ch
ange
s bi
odiv
ersit
y an
d he
tero
gene
ity,
both
abo
ve-
stra
ins
of a
var
iety
of g
ener
a of
Pro
teob
acte
ria in
clud
ing
Rhizo
bium
of p
heno
lics
and
aldo
nic
acid
s fr
om r
oots
, is
of v
ital i
mpo
rtan
ce fo
r at
mos
pher
ic d
initr
ogen
(N2
the
tree
and
shr
ub g
ener
a Al
nus,
Casu
arin
a, C
eano
thus
, and
Myr
ica, a
tmos
pher
ic N
2 is s
uppl
ied
by
Anab
aena
pro
vide
s at
mos
pher
ic N
to
wat
erfe
rns (
Azol
laas
100
kg
N h
a-1
55
4
cyan
obac
teria
und
er h
igh
P lo
ads
(Lam
ers
et a
l
gas (
Knei
p et
al.,
200
7; K
hade
m e
t al.,
201
0; R
eed
et a
l ro
ots (
Hodg
e et
al
Be
low
grou
nd ca
rbon
turn
over
Spha
gnum
1995
; Haj
ek e
t al
Spha
gnum
can
, for
inst
ance
, not
56
Spha
gnum
pea
t and
conc
omita
ntly
et
al
et a
l
Aste
lia p
umila
and
et a
l
et a
lPs
eudo
mon
as c
hlor
orap
his
an
d (B
odel
ier e
t al
3- an
d O
H-
acce
ptor
, and
pH
in th
e rh
izosp
here
. As a
resu
lt of
RO
L or
tem
pora
ry w
ater
tabl
e dr
aw-d
own
(e.g
.,
of a
mm
oniu
m a
nd n
itrite
by
nitr
ifyin
g pr
oteo
bact
eria
(e.g
., th
e ge
nera
,
, 42-
prot
eoba
cter
ia (e
.g.,
, Thi
obac
illus
Acid
ithio
bacil
lus
,
is st
rong
ly re
gula
ted
by p
H. T
he p
H in
the
rhizo
sphe
re m
ay b
e m
uch
low
er th
an th
at o
f the
bul
k 57
4
prot
on lo
ss b
y ro
ots (
e.g.
, for
NH 4+
+
arou
nd p
H 6-
6.5
by th
e CO
2-HCO
3-
HCO
3-
soils
(Van
der
Hei
de e
t al
N
itrog
en: a
mob
ile n
utrie
nt th
at m
ay b
ecom
e to
xic
Eith
er th
e av
aila
bilit
y of
ele
ctro
n do
nors
or
the
avai
labi
lity
of e
lect
ron
acce
ptor
s de
term
ines
the
bulk
soil.
Am
mon
ium
(NH 4+
3-2-
et a
l
3-4+
4+ 4+ c
an
4+ or
NO
3- , dep
endi
ng o
n th
e pH
of t
he s
urfa
ce w
ater
whe
re th
e sp
ecie
s m
ainl
y oc
curs
(Roe
lofs
et a
l4+ u
ptak
e, a
lthou
gh N
H 4+
3- . Pla
nts o
f rip
aria
n w
etla
nds,
how
ever
, are
kno
wn
to fa
ce p
robl
ems a
t a
high
NH 4+ :
NO
3-et
al
., 20
03; V
an d
en B
erg
et a
l4+ in
puts
, as a
resu
lt of
am
mon
ia
(NH 3
et a
l4+ :
NO
3-
4+
(NH 3
3et
al
et a
l It
has
been
sho
wn
that
incr
ease
d av
aila
bilit
y of
NO
3-2-
58
et a
l., 1
989;
Art
h et
al et
al.,
199
8; B
urgi
n an
d Ha
milt
on, 2
007;
Hu
et a
lTh
e up
take
of N
H 4+ or N
O3-
mar
ine
wet
land
s by
redu
cing
the
N su
pply
(Bod
elie
r et a
l., 1
996;
Wel
sh e
t al
2 and
N2O
2
2 2O
:N2
2O
et a
l2O
em
issio
n
et a
l
et a
l Hi
gh in
puts
of N
O3-
et a
lrh
izosp
here
(Luc
asse
n et
al
2+
3+
3- in o
rder
to c
hang
e
arch
aea
conv
erts
ferr
ic co
mpo
unds
into
ferr
ous
iron
(Lov
ley
& P
hilli
ps, 1
988;
Col
eman
et a
l., 1
993;
Lo
vley
et a
l., 2
004;
Wei
ss e
t al
et a
l
et a
l
et a
l., 1
984;
Sm
olde
rs &
Roe
lofs
, 199
6; E
mer
son
et a
l
59
4
et a
l
Raba
ey e
t al
both
Bac
teria
(e.
g.,
, Ar
chae
oglo
bus
2S
(SO
42-2
othe
r met
al co
ntai
ning
enz
ymes
(Koc
h et
al.,
199
0; R
aven
& S
crim
geou
r,
2
2S, H
S- and
S2-
, pK 1
2
H 2S w
ill b
e pr
evai
ling
due
to H
CO3—
CO2
2
is kn
own
to h
ampe
r th
e es
tabl
ishm
ent
and
grow
th o
f th
e tw
o m
ain
man
grov
e ge
nera
Avi
cenn
ia a
nd R
hizo
phor
ase
agra
sses
incl
udin
g Th
alas
sia te
stud
inum
and
Zos
tera
mar
ina
(Koc
h et
al
et a
l., 2
008;
Hol
mer
et a
lRO
L of
sea
gras
s pl
ants
lead
to lo
w H
2
grow
ing
seas
on d
ecre
asin
g RO
L, h
ighe
r m
icro
bial
O2
and
H 2w
etla
nds (
Azzo
ni e
t al
wat
er h
ave
raise
d SO
42-
fresh
wat
er w
etla
nd s
oils,
SO
42-
et a
lca
n be
det
erm
ined
by
the
avai
labi
lity
of e
ither
ele
ctro
n do
nors
or S
O42-
(L
amer
s et a
lL-1
et a
l.,
1968
; Arm
stro
ng e
t al.,
199
6; S
mol
ders
and
Roe
lofs
, 199
6; L
amer
s et a
l.,
1998
a; G
eurt
s et a
l2
60
adde
d H
S in
an
anae
robi
c rh
izotr
on w
ith li
ving
pla
nts
and
natu
ral o
rgan
ic w
etla
nd
and
S
prec
ipita
tes h
ave
been
Lam
ers &
Jans
sen,
unp
ublis
hed
resu
lts.
2
2re
leas
ed t
o th
e at
mos
pher
e (L
oman
s et
al
et a
let
al
42- w
ill s
erve
as
a te
rmin
al e
lect
ron
acce
ptor
, it
will
spe
ed u
p 42-
2et
al
., 19
89; R
oden
& E
dmon
ds, 1
997;
Lam
ers
et a
l., 1
998a
; Sm
olde
rs e
t al.,
et a
l., 2
002a
; Loe
b et
al.,
200
8; G
eurt
s et
al
in O
2
61
4
the
soil
can
be u
sed
as a
n el
ectr
on d
onor
by
prot
eoba
cter
ia su
ch a
s Thi
obac
illus
coup
led
to t
he u
se o
f NO
3- (Haa
ijer
et a
lor
gani
c co
mpo
unds
, NO
3-
wet
land
s whe
re th
is po
llute
d w
ater
is d
ischa
rged
. In
this
way
, the
re is
a m
icrob
ial ‘
S br
idge
’ bet
wee
n N
an
d P
at th
is hy
drol
ogica
l sca
le (S
mol
ders
et a
l2
0 and
2
et a
l
As a
res
ult
of a
nthr
opog
enic
forc
ing,
the
inpu
t of
all
thre
e el
emen
ts, N
, S a
nd P
, int
o w
etla
nds
has
stro
ngly
cha
nged
at
a gl
obal
sca
le (
Vito
usek
et
al
pollu
ted
by N
and
P (C
arpe
nter
et a
l a
nd N
H y
in re
gion
s with
inte
nsiv
e fa
rmin
g (V
itous
ek e
t al.,
199
7; B
obbi
nk e
t al
em
issio
ns
use
of c
oal a
s a fu
el (S
hah
et a
ldr
aine
d fo
r la
nd u
se c
hang
e (L
amer
s et
al
wet
land
s ha
s be
en c
hang
ed (L
amer
s et
al
biod
iver
sity
have
bee
n al
tere
d. T
he re
silie
nce
of e
cosy
stem
s to
glo
bal c
hang
e de
pend
s to
a la
rge
inco
rpor
ated
in e
cosy
stem
mod
els (
Tres
eder
et a
l
6263
4
6465
Gene
ral c
oncl
usio
n
rhizo
sphe
re re
sear
ch, i
n w
hich
bas
ic an
d ap
plie
d ec
olog
ical i
ssue
s on
stud
ents
shou
ld b
e ch
alle
nged
and
trai
ned
to a
ddre
ss e
colo
gica
l top
ics
it is
vita
l tha
t bot
h m
icrob
iolo
gist
s and
pla
nt e
colo
gist
s mee
t in
the
4
m
anag
emen
t and
exp
erim
enta
l lim
ing
Ab
stra
ctRi
ch fe
ns a
re k
now
n fo
r th
eir
high
bot
anic
al d
iver
sity
enco
mpa
ssin
g m
any
enda
nger
ed s
peci
es.
to m
aint
ain
a hi
gh b
iodi
vers
ity b
y m
eans
of s
low
ing
dow
n th
e na
tura
l suc
cess
ion
from
cal
care
ous
cove
r do
min
ated
by
Spha
gnum
esta
blish
men
t an
d do
min
ance
of S
phag
num
spp
. ser
ious
ly c
ompl
icat
ed t
he d
evel
opm
ent
of r
ich 67
spec
ies
(Boo
tsm
a et
al.,
the
peat
mat
. Thi
s pr
oces
s cr
eate
s a
biog
eoch
emic
al g
radi
ent t
hat e
nhan
ces
the
deve
lopm
ent o
f
fen
(with
or
with
out
hum
moc
k fo
rmin
g Sp
hagn
um
(Ver
hoev
en &
Bob
bink
, 200
1; G
root
jans
et
al.,
than
poo
r fen
s or w
oodl
ands
. The
se e
arlie
r suc
cess
iona
l sta
ges a
re, h
owev
er, s
erio
usly
thre
aten
ed
and
biod
iver
sity
has s
tron
gly
decl
ined
in m
any
rich
fens
as a
resu
lt of
fast
succ
essio
n to
eith
er p
oor
fens
or w
oodl
ands
(Bel
tman
et a
l., 2
001;
Mid
dlet
on e
t al.,
200
6a,b
; Lam
ers e
t al
(Hog
g et
al.,
199
5; K
ooijm
an &
Pau
lisse
n, 2
006;
Klim
kow
ska
et a
l., 2
007;
Lam
ers
et a
l., 2
002b
;
Spha
gnum
dom
inat
ed p
oor f
ens (
van
Digg
elen
et
al.,
1996
; Koo
ijman
& P
aulis
sen,
200
6; B
obbi
nk e
t al
., 20
10; L
amer
s et
al
Spha
gnum
(Hem
ond,
198
0; V
erho
even
& L
iefv
eld,
199
7; B
oots
ma
et a
l.,
et a
l., 1
995;
Bob
bink
et a
l., 1
998;
Bob
bink
et a
l., 2
010;
Cus
ell e
t al
graz
ing
or b
urni
ng (H
ogg
et a
l., 1
995;
Mid
dlet
on e
t al.,
disp
erse
or g
erm
inat
e (M
iddl
eton
et a
let
al.,
199
6;
van
Belle
et a
l
et a
l.,
Spha
gnum
(Bel
tman
et a
l m
easu
re to
cou
nter
act
Spha
gnum
spp.
from
bec
omin
g do
min
ant (
Beltm
an e
t al.,
200
1; D
orla
nd
68
et a
lsp
ecie
s to
re-
esta
blish
(Be
ltman
et
al.,
1996
a; 2
001;
Pat
zelt
et a
l
1991
; Sm
olde
rs e
t al Sp
hagn
um
M
ater
ial a
nd M
etho
ds
Rese
arch
are
aTh
e re
sear
ch w
as c
arrie
d ou
t in
the
wet
land
res
erve
and
Nat
ura
2000
are
a th
e ‘N
ieuw
koop
se
rese
rve,
as
wel
l as
seve
ral l
arge
sha
llow
fen
lake
s (d
en H
eld
et a
l
by h
uman
inte
rfere
nce,
as
area
s w
ithou
t m
anag
emen
t ha
ve a
ll de
velo
ped
into
car
r w
oodl
and
1.5
met
res
belo
w s
ea l
evel
, w
hich
is
0.4
to 4
met
res
high
er t
han
surr
ound
ing
agric
ultu
ral a
reas
tha
t ha
ve s
ubsid
ed d
ue t
o dr
aina
ge. A
s a
resu
lt, t
he
high
surf
ace
wat
er le
vels.
sele
cted
plo
ts th
roug
hout
the
rese
rve.
In 2
013,
49
of th
ese
hist
oric
fen
sites
wer
e re
asse
ssed
. The
2 to
30
m2 (a
vera
ge si
ze o
f 10
m2
had
been
car
ried
out a
nnua
lly fo
r at l
east
10
year
s bef
ore
the
star
t of t
his p
art o
f the
rese
arch
.
indi
vidu
al, 2
: 2-5
indi
vidu
als;
3: 6
-50
indi
vidu
als,
4: >
50 in
divi
dual
s; 5
: 5-1
2% co
ver;
6: 1
3-25
% co
ver;
7: 2
6-50
% c
over
; 8: 5
1-75
% c
over
; 9: 7
6-10
0% c
over
. The
num
bers
of i
ndiv
idua
ls fo
r cat
egor
ies
1-4
fact
or ca
lcula
ted
as th
e lo
g of
the
area
of t
he re
levé
, div
ided
by
the
log
of th
e st
anda
rd re
levé
are
a (4
m
2
69
for b
ryop
hyte
s, bu
t an
d Sp
hagn
um ru
bellu
m w
ere
cons
ider
ed o
ne s
pecie
s.
Li
min
g ex
perim
ent
Spha
gnum
3 84%
, MgC
O3
of n
itrat
e (N
O3-
4++
calc
ium
(Ca2+
2+
2
clos
ed c
ham
ber c
onta
inin
g 0.
2 M
H3PO
42
22
2 and
HCO
3-
ml 6
5% H
NO
3-
2+, M
g2+
3- (Kam
phak
e et
al
4+
phos
phat
e (P
O43-
+
70
for b
ryop
hyte
s an
d va
scul
ar p
lant
s se
para
tely,
usin
g CA
NO
CO (V
ersio
n 4.
55; t
er B
raak
& Š
mila
uer,
resid
uals
wer
e no
t nor
mal
ly d
istrib
uted
.
Re
sults
Hi
stor
ical
refe
renc
es
Spha
gnum
of p
oor f
ens.
In 4
6 of
the
hist
oric
rele
vés S
phag
num
pal
ustr
e or
w
as th
e do
min
ant
Pelli
a (c
ode
. Vas
cula
r pla
nts o
f ric
h fe
ns su
ch a
s Ham
mar
bya
palu
dosa
and
Lip
aris
loes
elii
wer
e pr
esen
t
71
Palla
vicin
ia ly
ellii
a
nd
vasc
ular
pla
nt s
pecie
s su
ch a
s Dr
yopt
eris
cart
husia
na
, Suc
cisa
prat
ensis
,
such
as S
phag
num
mag
ella
nicu
mS.
rube
llum
Spha
gnum
pa
lust
re
spec
ies S
. mag
ella
nicu
m
spec
ies C
epha
lozia
mac
rost
achy
a Sp
hagn
um sp
ecie
s.
72
, S. p
apill
osum
a
nd S
. rub
ellu
m
eac
h
cont
aine
d ra
re a
nd re
d-lis
t spe
cies
, the
bur
ned
rele
vés
gene
rally
onl
y co
ntai
ned
com
mon
spe
cies
.
S. p
alus
tre
S.
,
. In
only
3 re
levé
s a c
lose
d br
yoph
yte
cove
r was
pre
sent
as P
hrag
mite
s aus
tral
isM
olin
ia ca
erul
ea, P
euce
danu
m
palu
stre
, Jun
cus
subn
odul
osus
(8
.
2 ye
ars,
whi
le fo
r the
oth
er lim
e tr
eatm
ents
pH
grad
ually
dec
reas
ed a
nd re
ache
d in
term
edia
te va
lues
.
73
trea
tmen
ts.
2+ a
nd M
g2+
2+ a
nd M
g2+
3-
betw
een
trea
tmen
ts w
ere
foun
d an
ymor
e. C
O2
4+ av
aila
ble
NO
3-
plan
t av
aila
ble
K+
43-, N
O3- , N
H 4+ and
K+
74
betw
een
trea
tmen
ts.
In a
ll lim
e tr
eatm
ents
, Sph
agnu
m
Spha
gnum
sta
rted
in a
ccor
danc
e w
ith
cove
r of
S. p
alus
tre
and
Pol
ytric
hum
com
mun
eth
e pr
esen
ce o
f ric
h fe
n sp
ecie
s lik
e An
eura
pin
guis
Lipa
ris lo
esel
iiP.
com
mun
ech
arac
teris
ed b
y S.
pal
ustr
eP.
com
mun
eS.
. I
n 3
of th
ese
rele
vés,
onl
y th
ese
thre
e br
yoph
yte
spec
ies w
ere
pres
ent. S.
pa
lust
re, S
. rub
ellu
m (1
8 C
. con
nive
ns
S. p
alus
tre
(11
S. ru
bellu
mC.
mac
rost
achy
a (4
rel
evés
; S.
pap
illos
um
S.
mag
ella
nicu
m
thro
ugho
ut 4
.5 y
ears
. How
ever
, one
1k
plot
, and
tw
o 2k
plo
ts w
ere
char
acte
rised
by
a ric
h fe
n
Spha
gnum
75
trea
tmen
ts.
Di
scus
sion
stro
ngly
det
erm
ined
by t
he d
evel
opm
ent o
f Sph
agnu
m co
ver. S
umm
er m
owin
g enh
ance
d Sp
hagn
um
avai
labi
lity
(Old
e Ve
nter
ink
et a
lPh
ragm
ites
aust
ralis
M
olin
ea c
aeru
lea
(Hog
g et
al.,
et a
l., 1
996;
Ko
tow
ski e
t al
., 20
01; M
iddl
eton
et
al.,
2006
a; G
root
jans
et
al
man
agem
ent m
easu
re se
ems p
erfe
ct to
dev
elop
spec
ies-
rich
poor
fens
with
Sph
agnu
m h
umm
ocks
,
76
Bu
rnin
g le
ads t
o gr
amin
oid
dom
inan
ce
Spha
gnum
gr
owth
. Sha
w e
t al.
on th
e lo
nger
term
esp
ecia
lly sh
adin
g by
vas
cula
r pla
nts (
Mid
dlet
on e
t al.,
with
a d
ecre
ased
bry
ophy
te c
over
indi
cate
s in
crea
sed
nutr
ient
ava
ilabi
lity
(Bob
bink
et a
l., 1
998;
M. c
aeru
lea
et a
l.,
Spha
gnum
Spha
gnum
cove
r and
enh
ance
d th
e
et a
lSp
hagn
um h
ad b
een
Spha
gnum
and
esta
blish
men
t of S
phag
num
(Bel
tman
et a
l.,
älso
n et
al.,
(Don
ath
et a
l., 2
007;
van
Dur
en e
t al
., 19
98; B
eltm
an e
t al
.,
as h
ydro
chor
y is a
n im
port
ant d
isper
sal m
echa
nism
in fe
ns (V
ogt e
t al.,
disp
ersa
l by s
prea
ding
pro
pagu
les (
Mäl
son
et a
l., 2
010;
Hed
berg
et a
l., 2
012;
Mid
dlet
on e
t al.,
et
al.,
4+ et
al
et a
lof
NH 4+
4+
4+
et a
l
77
spec
ies (
Boey
e et
al.,
199
7; B
obbi
nk et
al.,
199
8; O
lde
Vent
erin
k et a
l., 2
002;
Cus
ell e
t al.
acce
lera
te s
ucce
ssio
n to
war
ds S
phag
num
Spha
gnum
bec
ame
dom
inan
t aga
in in
et a
l., 1
994;
La
mer
s et a
let
al
Spha
gnum
spec
ies m
ay th
eref
ore
have
(Hog
g et
al.,
199
5; M
alm
er e
t al.,
200
5; K
ooijm
an &
Pau
lisse
n, 2
006;
Bob
bink
et a
l., 2
010;
Lam
ers
et a
let
al
et a
l., 1
996;
Koo
ijman
& P
aulis
sen,
200
6; B
obbi
nk e
t al
., 20
10; L
amer
s et a
let
al.,
199
4; V
erho
even
fen
stag
es o
n th
ick,
old
pea
t mat
s.
very
sho
rt te
rm, S
phag
num
Spha
gnum
may
hav
e be
en fa
vour
ed
Spha
gnum
had
not
bee
n re
mov
ed b
efor
e
of S
phag
num
et a
l., 2
009;
Lam
ers
et a
l
mow
ing
regi
me.
78
Spha
gnum
spec
ies n
ot n
eces
saril
y inh
ibit
the
occu
rren
ce
Spha
gnum
(van
Dig
gele
n et
al.,
199
6; B
eltm
an e
t al.,
200
1; L
amer
s et
al.,
et a
lw
inte
r mow
ing,
and
lim
ing
wer
e al
l una
ble
to re
stor
e ric
h fe
ns in
the
long
term
.
79
74
6 |
How
to c
hoos
e be
twee
n ev
ils:
Th
e bi
ogeo
chem
istry
of d
rain
ed co
asta
l
peat
land
s cau
ses a
man
agem
ent d
ilem
ma
Jo
seph
a M
.H. v
an D
igge
len,
Leo
n P.
M. L
amer
s, Jo
hann
es H
.T. L
oerm
ans,
Ab
stra
ct
2 em
issio
n an
d se
vere
land
sub
siden
ce.
42-
42- to
adj
acen
t sur
face
wat
ers,
4
2
42-
man
agem
ent s
trate
gy in
dra
ined
coas
tal p
eatla
nds.
81
6
et a
l., 2
008;
Co
uwen
berg
et a
l., 2
010;
Koh
et a
l
et a
l
stro
ng C
sour
ces w
ith n
et e
miss
ion
rate
s ran
ging
bet
wee
n 80
and
880
g C
m-2 y
-1 (K
asim
ir-Kl
emed
tsso
n et
al.,
199
7; L
amer
s et
al
subs
iden
ce ra
tes
of 8
-10
mm
y-1 in
Wes
tern
Eur
ope
(Oud
e Es
sink
et a
l., 2
010;
Hoo
glan
d et
al
25-7
5 m
m y
-1 in
the
Unite
d St
ates
(Gal
low
ay e
t al
-1 in
Asia
hav
e be
en
obse
rved
(Syv
itski
et a
l
Mea
nwhi
le, t
he g
loba
l mea
n se
a le
vel is
risin
g at
an
incr
easin
g ra
te (I
PCC,
201
4; H
ay e
t al
Herb
ert e
t al.
et a
l., 2
004;
Zak
et a
l.,
2008
; Geu
rts e
t al
the
tops
oil,
and
leac
hing
of g
roun
dwat
er fr
om th
e re
duce
d su
bsoi
l (va
n Be
ek e
t al
82
incr
ease
d su
lpha
te (S
O42-
SO42-
et a
l., 1
998a
; 200
2a; S
mol
ders
et
al.,
peat
land
s (Jo
oste
n et
al.,
201
2; G
root
jans
et a
l2
2
et a
lm
ay a
lso le
ad t
o hi
gh m
etha
ne (C
H 4to
tal G
HG e
miss
ion
(e.g
. Moo
re &
Kno
wle
s, 19
89; H
endr
iks
et a
l
wat
er (A
ldou
s et
al.,
200
7; Z
ak e
t al.,
201
0; C
abez
as e
t al.,
201
3; V
an d
e Ri
et e
t al.,
201
3; K
insm
an-
Cost
ello
et a
l
for w
ater
man
agem
ent,
natu
re m
anag
emen
t and
pol
icy w
ith re
spec
t to
larg
e co
asta
l pea
tland
are
as.
and
disc
uss t
he e
mer
ging
dile
mm
as in
term
s of p
refe
rred
wat
er m
anag
emen
t and
futu
re p
olicy
for a
m
ore
sust
aina
ble
use
of su
bsid
ing
coas
tal p
eatla
nd re
gion
s.
2 whe
n th
e gr
ound
wat
er le
vel d
rops
. Eve
n w
ithou
t
& C
lark
e, 2
002;
Joos
ten
et a
l2
to a
net
rele
ase
of C
O2 an
d N
2O (K
asim
ir-Kl
emed
tsso
n et
al
83
6
2 em
issio
n ra
tes,
how
ever
, aer
obic
ra
tes s
how
a 2
-fold
incr
ease
in th
e un
satu
rate
d pe
at zo
ne w
hen
com
pare
d to
the
satu
rate
d pe
at zo
ne
et a
let
al
et a
l
3- et
al.,
et a
l
et a
l., 2
010;
Agg
enba
ch e
t al
., a
nd o
rgan
ic S
pres
ent i
n
2+ a
nd S
O42-
(Sm
olde
rs e
t al.,
200
6b; H
aaije
r et a
l42-
eas
ily
not a
ccum
ulat
e in
this
zone
and
the
tota
l S p
ool i
s ge
nera
lly o
nly
sligh
tly h
ighe
r in
the
unsa
tura
ted
2+
84
day
4 em
issio
n ra
tes w
ere
negl
igib
le in
et a
l, 20
05;
et a
l
et a
l., 1
994;
van
Bee
k et
al
85
6
42-
et a
lw
hile
SO42-
the
unsa
tura
ted
peat
zon
e (v
an B
eek
et a
l42-
will
tem
pora
rily
lead
to a
stro
ng in
crea
se o
f its
42- is
2bo
unda
ry, in
tern
ally
mob
ilisin
g pho
spha
te (P
O43-
2et
al.,
198
9; S
mol
ders
& R
oelo
fs, 1
995;
Sm
olde
rs e
t al.,
200
6a; L
amer
s et a
l
rela
ted
dom
inan
ce o
f fas
t gro
win
g prim
ary p
rodu
cers
such
as d
uckw
eed
or al
gae
(Roe
lofs
, 199
1; C
orel
l,
42-
86
4
4
satu
rate
d pe
at zo
ne.
2Co
uwen
berg
et a
l., 2
010;
Joos
ten
et a
l., 2
012;
Gro
otja
ns e
t al
4 em
issio
ns
et a
l.,
2007
; Har
pens
lage
r et a
l., su
bm.
et a
l., 2
013;
Hen
driks
et a
l., 2
007;
Hen
driks
et a
l42-
4et
al
42-
wet
pea
tland
s. As
a re
sult,
gro
undw
ater
SO
42-
2+
87
44
6
stro
ngly
mob
ilise
d to
the
pore
wat
er a
nd g
roun
dwat
er (A
ldou
s et
al.,
200
7; Z
ak e
t al.,
200
8; 2
010;
Ca
beza
s et a
l., 2
013;
van
de
Riet
et a
lof
PO
43-
43-et
al.,
199
8a; C
abez
as e
t al
., 20
13; K
insm
an-C
oste
llo e
t al
Dijk
et a
lLu
cass
en e
t al.,
200
5; v
an d
e Ri
et e
t al.,
201
3; K
insm
an-C
oste
llo e
t al
et
al.,
2004
; Za
k et
al.,
201
0; K
insm
an-C
oste
llo e
t al
drai
nage
thr
ough
sha
llow
pat
hway
s, e
nhan
cing
nut
rient
tra
nspo
rt f
rom
the
form
er u
nsat
urat
ed
et a
l
of n
utrie
nts.
In
gene
ral,
grou
ndw
ater
mov
emen
t w
ill b
e lim
ited
com
pare
d to
dra
inag
e-ba
sed
will
occ
ur p
redo
min
antly
dur
ing
heav
y ra
infa
ll.
2+
upo
n hi
gh w
ater
leve
ls in
duce
s co
-
3+ (S
mol
ders
et a
l., 2
006a
; Geu
rts
et a
l., 2
008;
Lam
ers
et a
l.,
88
Th
e ch
oice
bet
wee
n ev
ils: r
aisi
ng th
e gr
ound
wat
er le
vel
P in
the
uns
atur
ated
zon
e of
coa
stal
pea
tland
s th
at, u
nlike
cha
nges
in G
HG e
miss
ions
(M
oore
&
Know
les,
1989
; Kas
imir-
Klem
edts
son
et a
l., 1
997;
Hen
drik
s et a
l
episo
des o
f hig
h SO
42-
2S (L
amer
s et
al
Beek
et a
l., 2
004;
200
7; H
endr
iks e
t al
mov
emen
t in
the
satu
rate
d an
d un
satu
rate
d zo
ne s
houl
d al
so b
e ta
ken
into
acc
ount
to d
eter
min
e
chan
ge a
nd s
ea le
vel r
ise, c
urre
nt d
rain
age-
base
d la
nd u
se a
nd w
ater
man
agem
ent
is co
nsid
ered
un
sust
aina
ble,
not
onl
y w
ith re
spec
t to
GHG
emiss
ions
and
soil
subs
iden
ce, b
ut a
lso w
ith re
spec
t to
2+ w
ith P
, but
also
bec
ause
of m
ore
limite
d w
ater
mov
emen
t tow
ards
the
surfa
ce w
ater
89
6
(Ald
ous e
t al.,
200
7; K
insm
an-C
oste
llo e
t al
et a
l
et a
l
to a
djac
ent s
urfa
ce w
ater
s. O
f cou
rse
the
sugg
este
d ch
ange
s nee
d su
ppor
t of l
and
owne
rs a
nd o
ther
et a
l., 2
007;
Em
sens
et a
l.,
et a
let
al
90
Conc
lusi
ons
seve
re S
O4
mos
t sus
tain
able
wat
er m
anag
emen
t str
ateg
y in
the
long
term
.
91
6
7 |
Gen
eral
dis
cuss
ion
and
synt
hesi
s
Jose
pha
M.H
. van
Dig
gele
n
Peat
land
s and
thei
r lan
d us
e
thei
r sus
tain
able
futu
re m
anag
emen
t. W
orld
wid
e, th
e an
thro
poge
nic f
orcin
g on
wet
land
eco
syst
ems i
s
et a
l., 1
996;
Koe
rsel
man
& V
erho
even
, 199
2;
Joos
ten
& C
lark
e, 2
002;
Lam
ers e
t al
surr
ound
ings
. Now
aday
s a la
rge
part
of t
he te
rres
tria
l pea
t are
as in
the
Neth
erla
nds a
re p
redo
min
antly
Digg
elen
, 199
5; La
mer
s et a
l
93
A
B C
7
2
Drai
nage
-bas
ed la
nd u
se s
ever
ely
chan
ges
loca
l and
reg
iona
l hyd
rolo
gy a
nd li
mits
the
sup
ply
of
natu
ral s
yste
ms (
Boot
sma
et a
l., 2
002;
Cus
ell e
t al.
et a
l., 1
990;
van
Dig
gele
n et
al
enha
nced
Spha
gnum
Joos
ten
& C
lark
e, 2
002;
Sle
utel
et a
l., 2
003;
Bel
lam
y et
al.,
200
5; L
aiho
, 200
6; L
amer
s et
al
2et
al.,
201
2; G
root
jans
et
al
4 em
issio
n (C
hapt
er 6
; Zak
et
al.,
2010
; Har
pens
lage
r et a
l
2 ava
ilabi
lity
play
s a
very
impo
rtan
t ro
le (C
hapt
er 2
, 4 a
nd 6
; Sm
olde
rs e
t al
., 20
01; L
amer
s et
al.,
200
2a; H
upfe
r &
Le
wan
dow
ski,
2008
; Geu
rts e
t al
2 an
d hi
gh S
O42-
com
poun
ds a
t a m
illim
etre
94
2, hig
h N
O3-
42-2+
c
ompo
unds
in th
e pe
at s
oil (
Haai
jer e
t al.,
200
7;
Burg
in &
Ham
ilton
, 200
8; S
mol
ders
et a
l-ri
ch se
dim
ents
typi
cal
et a
l., 2
006a
; Geu
rts e
t al
surfa
ce w
ater
SO
42-et
al
.,42-
42- ri
ch
2
et a
l3-
et
alet
al
4+
et a
l.,
2 in a
naer
obic
wet
land
soils
, NO
3- is th
e m
ost f
avou
rabl
e el
ectr
on a
ccep
tor
2.
95
7
grow
ing
seas
on, r
ates
of m
icrob
ially
-med
iate
d NO
3-
enha
nced
SO
42-
et a
l.,
42- a
nd N
O3-
42-
and
NO3-
inpu
t of P
and
SO
42-et
al
-1 to
obt
ain
wel
l-dev
elop
ed s
ubm
erge
d et
al
by d
irect
ly in
crea
sing
O2
mac
roph
ytes
is in
hibi
ted
(e.g
. Bar
ko e
t al
96
4, S
O4
34
-1
coas
tal p
eatla
nd (v
an D
igge
len
et a
lei
ther
favo
urab
le C
hara
Char
a sp
p. ar
e
et a
l
Old
e Ve
nter
ink
et a
l., 2
002;
Else
r et
al
et
al (Bob
bink
et a
l., 1
998;
Old
e Ve
nter
ink,
200
2; C
usel
l et a
l
gram
inoi
d sp
ecie
s a
nd
97
AB
C
7
4+
43-et
al
., 20
13; K
insm
an-C
oste
llo e
t al
enha
nced
NH 4+
and
in
deed
sho
wed
net
CO
2ba
re p
eat
soils
sho
wed
CO
24+
et
al4+
4+
et a
l., 2
007;
Zhu
et a
lIn
the
rhizo
sphe
re o
f
and
aero
bic
CH4
et a
l.,
2 em
issio
n fro
m ro
ot
98
4
may
be c
onsid
ered
unf
avou
rabl
e by s
ome s
take
hold
ers,
as cu
rren
t dra
inag
e-ba
sed
wat
er m
anag
emen
t
long
er te
rm a
re h
igh
(Cos
tanz
a et
al
99
4
7
scen
ario
s will
be
disc
usse
d.
M
anag
emen
t sce
nario
s
Digg
elen
et a
l., 1
996;
Lam
ers
et a
l
et a
l
et a
l., 2
005;
Cus
ell e
t al
et a
l.,
(Boo
tsm
a et
al.,
200
2; C
usel
l et a
l.
100
Ag
ricul
tura
l are
as
of h
igh
SO42-
42-in
duce
d
et a
l
land
subs
iden
ce, w
hich
mig
ht n
ot o
nly
be u
ndes
ired
from
an
envi
ronm
enta
l poi
nt o
f vie
w, b
ut a
lso
et
al Ther
efor
e, th
is ch
ange
in la
nd u
se n
eeds
the
supp
ort o
f lan
dow
ners
, whi
ch m
ay b
e a
delic
ate
issue
. To
Man
y ag
ricul
tura
lly u
sed
peat
land
are
as a
re c
urre
ntly
situ
ated
at
low
er le
vels
in t
he la
ndsc
ape
of g
roun
dwat
er in
the
se a
reas
unl
ike in
the
hig
her
situa
ted
natu
re a
reas
. As
drai
nage
cos
ts a
re
proc
ess (
Chap
ter 6
; Em
sens
et a
l., 2
015;
van
de
Riet
et a
l., 2
013;
Har
pens
lage
r et a
l
agric
ultu
ral a
rea.
101
7
Gene
ral c
oncl
usio
n
102
103
7
8 |
Refe
renc
es
O
ecol
ogia
84,
391
-397
.
va
scul
ar sy
stem
s. N
ew P
hyto
logi
st 1
34, 6
01-6
14.
of
rice
. Soi
l Bio
l. Bi
oche
m. 3
0, 5
09-5
15.
cy
clin
g in
the
rhizo
sphe
re se
dim
ents
of a
eut
roph
ic R
uppi
a ci
rrho
sa m
eado
w (V
alle
Sm
arla
cca,
N
ethe
rland
s. T
hesis
. Rijk
suni
vers
iteit
Gron
inge
n. K
luw
er A
cade
mic
Pub
lishe
rs.
la
ndsc
ape
with
a fu
ll ra
nge
of su
cces
siona
l sta
ges.
Eco
logi
cal E
ngin
eerin
g 7,
299
-313
.
105
8
an
d lim
ing.
Eco
logi
cal E
ngin
eerin
g 17
, 21-
31.
31
5 pp
.Bo
bbin
k, R
., Hi
cks,
K.,
Gallo
way
, J.,
Spra
nger
, T.,
Alke
mad
e, R
., As
hmor
e, M
., Bu
stam
ante
, M.,
Cind
erby
, S.,
so
ils a
nd se
dim
ents
.
Berli
n He
idel
berg
, Spr
inge
r-Ver
lag,
331
-362
.
Bode
lier,
P.L.
E., S
orre
ll, B
., Dr
ake,
H.L
., Kü
sel,
K., H
urek
, T.,
P., R
einh
old-
Hure
k, B
., Lo
vell,
C.,
Meg
onig
al,
th
e rh
izosp
here
of w
etla
nd p
lant
s. In
: Bob
bink
, R.,
Beltm
an, B
., Ve
rhoe
ven,
J.T.
A., W
higm
an,
Pl
ant S
oil 1
34, 1
89-2
07.
106
w
ater
inte
rface
. Hyd
robi
olog
ia 1
70, 2
29-2
44.
Sc
i. To
tal E
nviro
n. 3
77, 4
39-4
43.
4+ cyc
ling:
A c
ellu
lar
4+
15
9, 5
67-5
84.
3- -driv
en S
O42-
an
d S
cycl
ing.
Eco
syst
ems 1
1, 9
08-9
22.
3-4+
gr
owth
and
N re
mov
al a
bilit
y of
Aco
rus c
alam
us a
nd Ir
is ps
euda
coru
s in
a hy
drop
onic
syst
em.
Ch
apm
an &
Hal
l, Lo
ndon
, 229
-289
.
107
8
su
lfate
-red
ucin
g ba
cter
ia. N
atur
e 36
1, 4
36-4
38.
na
tura
l cap
ital.
Nat
ure
387,
253
-260
.
Pl
ant a
nd S
oil 3
84, 1
53-1
68.
en
viro
nmen
ts. P
lant
Soi
l 245
, 35-
47.
So
il Bi
ol. B
ioch
em. 3
3, 8
53-8
66.
ni
trat
e on
thre
e he
athl
and
spec
ies.
Pla
nt E
colo
gy 1
35, 1
85-1
96.
po
lyph
osph
ate
in A
cine
toba
cter
spp.
Bo
tany
, Utr
echt
Uni
vers
ity.
th
e N
ethe
rland
s.
108
Pl
ant a
nd S
oil 2
65, 2
67-2
77.
Else
r, J.J
., Br
acke
n, M
.E.S
., Cl
elan
d, E
.E.,
Grun
er, D
.S.,
Harp
ole,
W.S
., Hi
llebr
and,
H.,
Nga
i, J.T
., Se
ablo
om,
ci
rcum
neut
ral p
H. A
ppl.
Envi
ron.
Mic
rob.
12,
478
4-47
92.
fe
ns: C
an w
e fo
rce
an e
cosy
stem
rese
t? E
colo
gica
l Eng
inee
ring
77, 2
25-2
32.
W
etla
nds E
col M
anag
e, 1
7, 7
1-84
.
Sc
ienc
e 31
9, 1
235-
1238
.
Li
mno
l. O
cean
ogr.
51, 2
308-
2318
.
lo
cks u
p ca
rbon
in p
eatla
nds b
y re
stra
inin
g a
singl
e en
zym
e. N
atur
e 40
9, 1
49-1
49.
109
8
cu
shio
n pl
ants
in P
atag
onia
. New
Phy
tolo
gist
190
, 398
-408
.
of
org
anic
soils
. Geo
logi
cal S
urve
y, Ci
rcul
ar 1
182.
su
ppre
ssio
n of
the
wet
land
met
hane
sour
ce in
the
20th
and
21s
t cen
turie
s.
w
ater
in fe
ns. W
ater
Res
earc
h 44
, 348
7-34
95.
th
e m
icro
envi
ronm
ent.
Soil
Biol
. Bio
chem
. 30,
190
3-19
16.
co
mpa
rison
with
oth
er m
etho
ds. H
ydro
biol
ogia
335
, 87-
95.
Hy
drob
iolo
gia
450,
99-
106.
W
iley,
Chic
hest
er, p
p 73
-90.
110
N
ew P
hyto
logi
st 1
64, 2
43-2
66.
Pl
ant e
colo
gy 1
76, 1
35-1
55.
Bi
ogeo
chem
istry
103
, 45-
57.
ba
lanc
e of
an
aban
done
d pe
at m
eado
w. B
ioge
osci
ence
s 4, 4
11-4
24.
a
grow
ing
thre
at to
fres
hwat
er w
etla
nds.
Eco
sphe
re.
Be
rkel
ey, 3
2 pp
.
111
8
ni
trog
en?
Tren
ds P
lant
Sci
. 5, 3
04-3
08.
N
ew P
hyto
logi
st 1
62, 9
-24.
Holm
er, M
., Pe
ders
en, O
., Kr
ause
-Jens
en, D
., O
lese
n, B
., Pe
ters
en, M
. H.,
Scho
pmey
er, S
., Ko
ch,
co
asta
l are
a. G
eode
rma
171-
172,
92-
97.
En
viro
nmen
tal T
echn
olog
y 20
, 727
-733
.Hu
, B.-I
., Ru
sh, D
., va
n de
r Bie
zen,
E.,
Zhen
g, P.
, Van
Mul
leko
m, M
., Sc
hout
en, S
., Da
mst
e, J.
S.S.
,
en
riche
d fr
om p
eat s
oil.
Appl
. Env
iron.
Mic
rob.
77,
966
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.
N
atur
e 40
2, 4
07-4
10.
He
lgol
ände
r Mee
resu
nter
s 49:
223
-236
.
Plan
t Bio
l. 1,
274
-287
.
Lo
gem
ann,
S.,
Muy
zer,
G., V
an L
oosd
rech
t, M
.C.M
., Ku
enen
, J.G
.
Pe
at S
ocie
ty, 3
04 p
p.
al
ong
an e
stua
rine
salin
ity g
radi
ent:
The
iron
conv
eyer
bel
t.
112
Ri
ce rh
izosp
here
. Sci
ence
195
, 179
-180
.
Gr
eenh
ouse
gas
em
issio
ns fr
om fa
rmed
org
anic
soils
: a re
view
. Soi
l Use
And
Man
agem
ent 1
3, 2
45-2
50.
m
etha
notr
oph
‘Met
hyla
cidi
philu
m fu
mar
iolic
um’ S
olV.
Mic
robi
olog
y-SG
M 1
56, 1
052-
1059
.
sy
mbi
osis.
BM
C Ev
ol. B
iol.7
, 55.
113
8
le
vel a
nd li
ght i
nten
sity.
Pla
nt E
colo
gy 1
55, p
. 147
-156
.
Lam
ers L
.P.M
., Vi
le, M
.A.,
Groo
tjans
, A.P
., Ac
rem
an, M
.C.,
Van
Digg
elen
, R.,
Evan
s, M
.G.,
Rich
ards
on,
Ca
mbr
idge
Uni
vers
ity P
ress
.
in
fres
hwat
er w
etla
nds.
Env
iron.
Sci
. Tec
hnol
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.
Gl
obal
Cha
nge
Biol
ogy
6, 5
83-5
86.
w
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Lim
nolo
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cean
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phy,
Vol.
47, N
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.
Lam
ers,
L.P
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Gove
rs, L
.L.,
Jans
sen,
I.C.
J.M.,
Geur
ts, J
.J.M
., va
n de
r Wel
le, M
.E.W
., va
n Ka
twijk
, M.M
.,
Lam
ers,
L.P
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Vile
, M.A
., Gr
ootja
ns, A
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Acre
man
, M.C
., va
n Di
ggel
en, R
., Ev
ans,
M.G
., Ri
char
dson
, C.J.
,
fe
ns in
Eur
ope
and
Nor
th A
mer
ica:
from
tria
l and
err
or to
an
evid
ence
-bas
ed a
ppro
ach.
Biol
ogic
al R
evie
ws,
1-2
2.
An
nals
of A
pplie
d Bi
olog
y 15
0, 1
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N
atur
e 44
2, 8
06-8
09.
114
co
mpo
unds
. Cel
l. M
ol. L
ife S
ci. 5
9, 5
75-5
88.
Go
rter
ia 7
, 102
-106
.
Ad
v. M
icro
bial
Phy
siol.
49, 2
19-2
86.
Luca
ssen
, E.C
.H.E
.T., B
obbi
nk, R
., Sm
olde
rs, A
.J.P.
, Van
der
Ven
, P.J.
M.,
Lam
ers,
L.P
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and
Roel
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67,
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.
va
scul
ar p
lant
s in
the
deve
lopm
ent o
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t-for
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g sy
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s.
J.
Ecol
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477
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.
So
il Bi
olog
y &
Bio
chem
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68,
317
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.
115
8
Ec
olog
ical S
tudi
es 1
91, 3
15 p
p.
M
ires a
nd P
eat,
Volu
me
1, 1
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Ge
olog
y 12
, 89-
103.
em
issio
ns fr
om p
eatla
nd so
ils. C
an. J
. Soi
l Sci
. 69,
33-
38.
pl
ants
in re
spon
se to
incr
ease
d ni
trog
en a
vaila
bilit
y. N
ew P
hyto
logi
st 1
26, 1
37-1
43.
co
ntro
ls of
subs
urfa
ce w
etla
nd b
ioge
oche
mist
ry. E
colo
gy 8
6, 3
334-
3344
.
ec
osys
tem
s in
Aust
ralia
. Aus
tral
ian
Jour
nal o
f Bot
any
51, 6
55-6
65.
W
aste
s Eng
inee
ring
33, 6
70-6
82.
Vom
Was
ser 2
1, 1
3-32
.
gr
ound
wat
er sy
stem
s: A
mod
elin
g st
udy
in th
e N
ethe
rland
s. W
ater
Res
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es R
esea
rch
46, 1
-16.
Ge
obot
any
18, p
p. 7
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116
Raba
ey, K
., Ro
drig
uez,
J., B
lack
all,
L. L
., Ke
ller,
J., G
ross
, P.,
Bats
tone
, D.,
Vers
trae
te, W
., an
d N
ealso
n, K
. H.
IS
ME
J. 1,
9-1
8.
N
atur
e 44
0, 9
18-9
21.
in
terfa
ce in
wet
land
s. L
imno
l. O
cean
ogr.
34, 1
004-
1013
.
Ann.
Rev
. Eco
l. Ev
ol. S
. 42,
489
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.
Co
mm
issie
inza
ke h
et zo
utge
halte
der
boe
zem
- en
pold
erw
ater
en v
an N
oord
-Hol
land
, ing
este
ld b
ij
Be
sluit
van
den
Min
ister
van
Wat
erst
aat v
an 2
4 ap
ril 1
939.
Rijk
suitg
ever
ij, ’s
-Gra
venh
age,
191
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ienc
e 22
8, 1
424-
1427
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e Kr
oon,
Be
rlin
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elbe
rg, S
prin
ger-V
erla
g, 1
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al
oide
s L. s
tand
s.
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idel
berg
, Ber
lin, S
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rum
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dem
ische
r Ver
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a
min
imal
mod
el. H
ydro
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l. 23
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83.
cu
cum
ber p
lant
s. P
lant
Soi
l 52,
403
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117
8
En
glish
Nat
ure
Rese
arch
Rep
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no.
172
.
Ca
rlier
, L.,
van
Cam
p, N
., Ve
rbee
ck, H
., va
n de
Wal
le, I
., Sa
mso
n, R
., Lu
st, N
., Le
meu
r,
du
e to
the
inle
t of r
iver
Rhi
ne w
ater
in p
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shal
low
wat
ers i
n Th
e N
ethe
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s.
Ar
chiv
f. H
ydro
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olde
rs,
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r Hyd
robi
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ie 1
50, 1
69-1
75.
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m p
hosp
hate
-enr
iched
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men
ts b
y ad
ding
vario
us ir
on co
mpo
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. Bio
geoc
hem
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54:
219
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.
Jo
urna
l of A
pplie
d Ec
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y 39
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Ge
oder
ma
132,
372
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.
th
e su
lphu
r brid
ge. B
ioge
oche
mist
ry 9
8, 1
-7.
118
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ure
181,
934
.
A.
W.,
Blee
ker,
A., V
erge
er, P
., Ar
roni
z-Cr
espo
, M.,
Lim
pens
, J.,
Lam
ers,
L.P
.M.,
Bobb
ink,
R.,
Dorla
nd,
Ch
apte
r 11
and
12. C
ambr
idge
Uni
vers
ity P
ress
.
Jo
urna
l of O
cean
ogra
phy
56, 6
67-6
73.
N
atur
e Ge
osci
ence
2, 6
81-6
86.
Am. J
. Bot
. 71,
666
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.CA
NO
CO R
efer
ence
man
ual a
nd u
ser’s
gui
de to
Can
oco
for W
indo
ws:
se
dim
ent,
Aarh
us B
ay, D
enm
ark.
Geo
chim
. Cos
moc
him
. Ac.
58,
511
5-51
29.
th
e un
-ioni
zed
amm
onia
spec
ies.
Env
iron.
Sci
. Tec
hnol
. 15,
837
-840
.
ph
osph
orus
, pot
assiu
m, a
nd li
me.
Soi
l Sci
ence
Soc
iety
of A
mer
ica
Jour
nal 4
2, 1
25-1
30.
pu
besc
ens a
nd M
olin
ia c
aeru
lea
on o
mbr
otro
phic
bog
s: ro
le o
f hig
h le
vels
of a
tmos
pher
ic n
itrog
en
Tres
eder
, K.K
., Ba
lser,
T.C.,
Brad
ford
, M.A
., Br
odie
, E.L
., Du
bins
ky, E
.A.,
Evin
er, V
.T., H
ofm
ocke
l, K.
S., L
enno
n,
N
atur
e 41
1, p
. 258
.
Van
Beek
, C.L
., D
roog
ers,
P., v
an H
arde
veld
, H.A
., va
n de
n Ee
rtw
egh,
G.A
.P.H
., Ve
lthof
, G.L
., O
enem
a,
119
8
Pl
ant P
hysio
logy
86,
914
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.
w
ith lo
w p
H. N
ew P
hyto
logi
st 1
66, 5
51-5
64.
Sc
i. To
tal E
nviro
n. 4
07, 2
325-
2332
.
pH
cau
ses h
igh
N2O
em
issio
ns in
a ri
paria
n ec
osys
tem
. Geo
biol
ogy
9, 2
94-3
00.
Van
der H
eide
, T.,
Smol
ders
, A.J.
P., R
ijken
s B.G
.A.,
van
Nes
, E.H
., va
n Ka
twijk
, M.M
., Ro
elof
s, J.
G.M
.
Em
piric
al e
vide
nce
from
por
e w
ater
ana
lyse
s. A
ppl.
Geoc
hem
. 25,
182
5-18
29.
Envi
ron.
Ei
ndra
ppor
tage
. Ond
erzo
ekce
ntru
m B
-WAR
E ra
ppor
tnr.
2013
.04,
264
pp.
Ne
derla
nd, b
edre
igin
g of
kans
?
H 2
120
Zo
ster
a m
arin
a. M
arin
e Ec
olog
y Pr
ogre
ss S
erie
s 157
, 159
-173
.
Sp
hagn
um. A
cta
Bota
nica
Nee
rland
ia 4
6, 1
17-1
30.
Ba
ckhu
ys P
ublis
hers
, Lei
den,
65-
87.
An
nals
of B
otan
y, 1-
9.
Vito
usek
, P.M
., Ab
er, J
.D.,
How
arth
, R.W
., Li
kens
, G.E
., M
atso
n, P.
A., S
chin
dler
, D.W
., Sc
hles
inge
r, W
.H.,
Ec
ol. A
ppl.
7, 7
37-7
50.
co
mpl
ete
subm
erge
nce.
New
Phy
tol.
170,
213
-226
.
m
eado
w. M
ar. E
col.-
Prog
r. Se
r. 20
8, 6
5-77
.
Bi
ogeo
chem
istry
102
, 135
-151
.
W
iley,
Chic
hest
er, 7
3-90
.
121
8
Eu
rope
an m
ires.
Ess
ay re
view
. Jou
rnal
of E
colo
gy 8
8, 1
87-2
03.
122
123
8
125
La
agve
enO
ver d
e he
le w
erel
d k
omen
‘wet
land
s’ vo
or in
zow
el zo
ut-,
brak
- en
zoet
wat
erge
bied
en. Z
e w
orde
n
afge
brok
en o
rgan
isch
mat
eria
al p
laat
svin
dt e
n ve
envo
rmin
g op
tree
dt. W
etla
nds
met
een
org
anisc
he
bode
m w
orde
n da
arom
vee
n- o
f moe
rasg
ebie
den
geno
emd,
die
doo
rgaa
ns o
nder
verd
eeld
wor
den
in o
mbr
otro
of h
oogv
een
en m
iner
otro
of la
agve
en. H
et v
ersc
hil h
iert
usse
n w
ordt
voo
ral b
epaa
ld d
oor
van
het w
ater
. Een
toen
ame
in d
e di
kte
van
het v
een
gaat
nam
elijk
gep
aard
met
een
afn
ame
van
gron
dwat
er-,
en e
en t
oena
me
van
rege
nwat
erin
vloe
d. H
ierd
oor
onts
taat
er
een
biog
eoch
emisc
he
2 ver
deel
d ov
er
men
sen
te v
inde
n om
en
nabi
j vee
ngeb
iede
n ov
eral
ter
wer
eld.
Om
dat
de m
ense
lijke
bev
olki
ng
nog
stee
ds g
roei
ende
is, n
eem
t de
druk
op
veen
gebi
eden
ook
nog
stee
ds to
e. G
rote
del
en v
an d
e ve
enge
bied
en zi
jn w
erel
dwijd
in g
ebru
ik g
enom
en te
n be
hoev
e va
n m
ense
lijke
voor
zieni
ngen
zoal
s
80%
in d
icht
bevo
lkte
del
en z
oals
Euro
pa, N
oord
-Am
erik
a en
Oos
t-Azie
, aan
vee
ngeb
ied
verlo
ren
9
In N
eder
land
is h
et la
agve
enla
ndsc
hap
het l
aags
t gel
egen
, en
kom
t voo
ral v
oor i
n he
t wes
telij
ke e
n
in d
e po
lder
s van
het
laag
veen
gebi
ed n
og st
eeds
vaa
k ee
n ty
pisc
h pa
troo
n te
zien
van
lang
gere
kte
en la
ter h
et a
grar
isch
gebr
uik
van
veen
wei
den
mog
elijk
te m
aken
was
stee
ds v
erde
re o
ntw
ater
ing
slote
n, k
anal
en, b
oeze
mw
ater
en e
n ge
mal
en in
en
rond
om d
e aa
ngre
nzen
de p
olde
rs is
ont
staa
n.
Door
win
dwer
king
en
golfs
lag
zijn
er u
itein
delij
k sm
alle
lega
kker
s w
egge
slage
n, w
aard
oor
ook
de t
erre
stris
che
dele
n va
n he
t la
ndsc
hap,
met
hie
rop
de v
eenw
eide
s di
e ve
elal
in
agra
risch
ge
brui
k zij
n, e
n ou
de l
egak
kers
of
veen
wei
des
met
blo
emrij
ke h
ooila
nden
, sc
hraa
lland
en e
n
De vo
ortd
uren
de d
rain
age
van
de la
agve
enge
bied
en, e
n oo
k van
om
ringe
nde
pold
ers,
hebb
en e
chte
r ge
leid
tot d
iver
se p
robl
emen
zoal
s ver
ande
ringe
n in
de
hydr
olog
ie (w
ater
teko
rt in
de
war
me
perio
den
als
agra
risch
e de
len
van
het
veen
gebi
ed. D
it ko
mt
door
dat
ze le
iden
tot
div
erse
pro
blem
en z
oals
er a
an d
e ha
nd va
n de
ze ke
nnis
een
door
kijk
gem
aakt
naa
r mog
elijk
hede
n vo
or d
uurz
aam
beh
eer,
om
de o
verg
eble
ven
laag
veen
gebi
eden
te b
ehou
den
of te
her
stel
len
in d
e to
ekom
st.
oppe
rvla
ktew
ater
met
3 v
ersc
hille
nde
salin
iteitn
ivea
us. E
r wer
d ve
rwac
ht d
at d
e ho
evee
lhei
d P
die
126
te z
ijn b
ij ee
n no
rmal
e of
lage
sal
inite
it. D
it ko
mt
zeer
waa
rsch
ijnlij
k do
orda
t de
top
laag
van
de
2+
2+
2+
de a
anw
ezig
e zu
urst
ofrij
ke t
opla
ag v
an d
e bo
dem
. Wan
neer
de
salin
iteit
in h
et o
pper
vlak
tew
ater
to
enee
mt,
neem
t oo
k su
lfaat
toe
wat
in d
e an
aero
be o
nder
wat
erbo
dem
wor
dt g
ered
ucee
rd t
ot
2+ in
de
topl
aag,
waa
rdoo
r er
min
der
P w
ordt
geb
onde
n en
dit
alsn
og n
aar h
et o
pper
vlak
tew
ater
verd
wijn
t. Ee
n ho
ge sa
linite
it do
et h
ierm
ee
4+
vert
oont
doo
rgaa
ns g
rote
ver
schi
llen.
Ond
anks
het
feit
dat p
lant
en N
H 4+ mak
kelij
k ku
nnen
opn
emen
4+
in h
un re
spon
s op
een
toen
ame
in N
H 4+
( kond
en g
roei
en b
ij ho
ge (2
mm
ol L
-1-1
4+
besc
hikb
aarh
eid
toen
eem
t in
wet
land
s, w
at ka
n le
iden
tot v
eran
derin
gen
in p
lant
ensa
men
stel
ling
ten
kost
e va
n de
bio
dive
rsite
it.
Beid
e so
orte
n lij
ken
op d
e ko
rte
term
ijn g
oed
best
and
tege
n ho
ge N
H 4+
hebb
en v
oor
hoge
NH 4+
aanp
asse
nd v
erm
ogen
van
dez
e so
ort a
fnee
mt b
ij aa
nhou
dend
hog
e N
H 4+
slech
ts 1
0% w
as b
ij Ve
ldru
s. De
ze v
ersc
hille
n in
gev
oelig
heid
kun
nen
mog
elijk
e de
ver
ande
ringe
n
NH4+
127
9
in ie
der g
eval
op
de k
orte
term
ijn, d
ie v
aak
leid
t tot
een
afn
ame
van
de w
etla
ndbi
odiv
ersit
eit b
ij ee
n to
enam
e va
n NH
4+ .
Vers
chill
ende
bio
geoc
hem
ische
pro
cess
en d
ie p
laat
svin
den
in d
e bo
dem
wor
den
gere
gule
erd
door
gew
orde
n. vo
edse
lbes
chik
baar
heid
voo
r pla
nten
. Dit
bepa
ald
in s
terk
e m
ate
de o
ntw
ikke
ling
en s
amen
stel
ling
gaan
ech
ter
hard
ach
teru
it in
Ned
erla
nd d
oor
de s
terk
e af
nam
e aa
n gr
ondw
ater
invl
oed
en h
et
gebr
ek a
an n
ieuw
e ve
rland
ing.
Div
erse
beh
eers
maa
treg
elen
, inc
lusie
f maa
ien
en b
rand
en, w
orde
n
128
mos
bede
kkin
g ge
dom
inee
rd d
oor
veen
mos
soor
ten
met
een
aan
tal r
ode-
lijst
soor
ten.
Bra
nden
(na
omst
andi
ghed
en v
aak
onom
keer
baar
ver
ande
rd z
ijn e
n hi
erdo
or d
e gr
ondw
ater
invl
oed
niet
te
Wer
eldw
ijd w
orde
n ve
enge
bied
en a
l zee
r la
ng g
edra
inee
rd o
m a
grar
isch
gebr
uik
te b
evor
dere
n,
2
aan
de k
ust v
aak
een
onge
kend
gro
te b
ron
van
sulfa
at v
oor a
angr
enze
nde
oppe
rvla
ktew
ater
en. D
it
agra
risch
e ve
enbo
dem
s wee
r ver
nat d
oor p
erm
anen
t hog
e gr
ondw
ater
peile
n in
te vo
eren
. Dit
leid
t ec
hter
vaa
k to
t een
toen
ame
van
met
haan
(CH 4
van
het v
oorm
alig
e CO
2
van
de v
eenw
eide
, wat
ver
volg
ens
kan
uits
poel
en n
aar
omlig
gend
e slo
ten
en d
aar
tot
dire
cte
en u
itsto
ot v
an b
roei
kasg
as,
zorg
t vo
or e
en s
erie
us d
ilem
ma
bij
wat
erbe
heer
ders
die
te
mak
en
veen
gebi
eden
in ku
stre
gio’
s. O
mda
t wat
erbe
heer
dire
ct is
gere
late
erd
aan
land
gebr
uik,
zou
de af
weg
ing
voor
een
nie
uw b
elei
d ni
et a
lleen
geb
asee
rd m
oete
n w
orde
n op
de
huid
ige
geoh
ydro
logi
sche
ligg
ing
129
9
van
natu
urlij
ke w
etla
nds,
en v
erw
acht
en w
e da
t een
per
man
ent h
oge
gron
dwat
erst
and
op d
e la
nge
zwav
elrij
ke v
eeng
ebie
den.
130
131
Tot s
lot
9
133
th, 1
982
in
Reek
, The
Net
herla
nds.
I liv
ed a
nd g
rew
up
in R
eek,
whe
re I
also
wen
t to
prim
ary
scho
ol.
Nijm
egen
. I a
m w
orki
ng a
s a co
nsul
tant
, and
mai
nly
focu
s on
appl
ied
ecol
ogica
l and
bio
geoc
hem
ical
rese
arch
in p
eatla
nd a
reas
thro
ugho
ut th
e Ne
ther
land
s. Se
vera
l pro
ject
s and
rese
arch
pro
gram
mes
liste
d be
low
. Jo
seph
a M
.H. v
an D
igge
len,
Leo
n P.
M. L
amer
s, Jo
hann
es H
.T. L
oerm
ans,
Alfo
ns J.
P. S
mol
ders
caus
es a
man
agem
ent d
ilem
ma.
Jose
pha
M.H
. van
Dig
gele
n, A
lfons
J.P.
Sm
olde
rs, E
ric J.
W. V
isser
, Séa
mas
Hic
ks, J
an G
.M. R
oelo
fs,
Jose
pha
M.H
. van
Dig
gele
n
Ecol
ogic
al E
ngin
eerin
g 75
, 208
-216
.
Jose
pha
M.H
. van
Dig
gele
n, L
eon
P.M
. Lam
ers,
Gijs
van
Dijk
, Maa
rten
J. S
chaa
fsm
a, Ja
n G.
M.
Lam
ers L
PM, V
an D
igge
len
JMH,
Op
den
Cam
p HJ
M, V
isser
EJW
, Luc
asse
n EC
HET,
Vile
MA,
Piet
er P
oot,
Step
hen
D. H
oppe
r, Jo
seph
a M
.H. v
an D
igge
len
Esth
er L
ucas
sen,
Ral
f Abe
n, A
lfons
Sm
olde
rs, R
olan
d Bo
bbin
k, Jo
se v
an D
igge
len,
Mic
hael
van
AJP
Smol
ders
, JM
H Va
n Di
ggel
en, J
JM G
eurt
s, M
DM P
oele
n, JG
M R
oelo
fs, E
CHET
Luc
asse
n,
José
van
Dig
gele
n
Alfo
ns S
mol
ders
, Jos
é va
n Di
ggel
en, J
ohan
Loe
rman
s, G
ijs v
an D
ijk, M
ark
van
Mul
leko
m, L
eon
Lam
ers L
PM A
JP S
mol
ders
, JM
H Va
n Di
ggel
en
veen
wei
den.
Tus
sen
Duin
en
Dijk
7: 3
0-36
.
Jose
pha
M.H
. Van
Dig
gele
n
The
6th
Jose
pha
M.H
. Van
Dig
gele
n, L
eon
P.M
. Lam
ers a
nd A
lfons
J.P.
Sm
olde
rs. C
hoos
ing
betw
een
Evils
:
day
2013
, Uni
vers
ity o
f Nijm
egen
, Net
herla
nds,
30
Oct
ober
201
3.
José
van
Dig
gele
nen
Pie
t-Jan
Wes
tend
orp.
Rui
m 4
jaar
ond
erzo
ek: D
uurz
aam
beh
eer W
orm
er- e
n Jis
perv
eld
(ora
l
23 se
ptem
ber 2
013.
Jose
pha
M.H
. Van
Dig
gele
n, A
lfons
J.P.
Sm
olde
rs a
nd L
eon
P.M
. Lam
ers.
Cho
osin
g be
twee
n Ev
ils:
th
Jose
pha
M.H
. van
Dig
gele
n, A
lfons
J.P.
Sm
olde
rs a
nd L
eon
P.M
. Lam
ers.
Cho
osin
g be
twee
n Ev
ils:
th
134
José
M.H
. van
Dig
gele
n, E
ric J.
W. V
isser
, Han
s de
Kroo
n, M
arku
s C. S
chm
id, H
uub
J.M. O
p de
n Ca
mp,
Jan
G.M
. Roe
lofs
and
Leo
n P.
M. L
amer
s.
th
José
M.H
. van
Dig
gele
n, E
ric J.
W. V
isser
, Han
s de
Kroo
n, M
arku
s Sch
mid
, Huu
b O
p de
n Ca
mp,
Jan
G.M
. Roe
lofs
and
Leo
n P.
M. L
amer
s.
3 De
cem
ber 2
009.
José
van
Dig
gele
n O
nder
zoek
naa
r een
duu
rzaa
m b
ehee
r van
het
W
orm
er- e
n Jis
perv
eld.
Ein
drap
port
age
2013
. In
opdr
acht
van
: Hoo
ghee
mra
adsc
hap
Holla
nds
Rap
port
num
mer
: 201
3.04
.
José
van
Dig
gele
n
Ond
erzo
ekce
ntru
m B
-WAR
E, N
ijmeg
en. R
appo
rtnu
mm
er: 2
013.
53.
Jero
en G
roen
endi
jk, T
om v
an d
en B
roek
, Ron
van
’t V
eer,
José
van
Dig
gele
n
José
van
Dig
gele
n he
t sed
imen
t in
de N
ieuw
koop
se p
lass
en. I
n op
drac
ht v
an: V
eren
igin
g N
atuu
rmon
umen
ten.
O
nder
zoek
cent
rum
B-W
ARE,
Nijm
egen
. Rap
port
num
mer
: 201
1.15
.
José
van
Dig
gele
n
B-W
ARE,
Nijm
egen
. Rap
port
num
mer
: 201
0.37
.
José
van
Dig
gele
n
In o
pdra
cht v
an: N
atuu
rmon
umen
ten.
Ond
erzo
ekce
ntru
m B
-WAR
E, N
ijmeg
en. R
appo
rtnu
mm
er:
2008
.45.
J.M.H
. van
Dig
gele
n
Nijm
egen
. Rap
port
num
mer
: 200
7.14
.
José
van
Dig
gele
nVe
enw
eide
gebi
ed H
DSR.
Ein
drap
port
age
nove
mbe
r 200
7. In
opd
rach
t van
: Hoo
ghee
mra
adsc
hap 13
5
136
137
Hier
zijn
we
dan
aan
het e
inde
gek
omen
. Nie
t alle
en h
et a
llerla
atst
e on
derd
eel o
m h
et g
ehee
l ec
ht a
f te
rond
en, m
aar h
et is
ove
r het
alg
emee
n oo
k no
g he
t mee
st g
elez
en h
oofd
stuk
in e
en
staa
n. S
inds
mijn
afs
tude
ren
in 2
007
ben
ik b
ij O
nder
zoek
cent
rum
B-W
ARE
in d
iens
t gek
omen
,
plek
ver
dien
en in
dit
dank
woo
rd, e
n da
t is n
aar m
ijn m
enin
g ei
genl
ijk a
an h
et e
inde
van
dit
kunn
en, m
et zo
wel
een
lach
als
een
traa
n. M
oni,
je b
ent e
en su
per l
ief,
moo
i en
gew
eldi
g m
ens!
Nu
zijn
er o
p on
ze k
anto
orka
mer
in to
taal
4 w
erkp
lekk
en, e
n de
laat
ste
wer
d ov
er d
e af
gelo
pen
jare
n af
wiss
elen
d be
zet d
oor J
ohan
Loe
rman
s en
Jero
en G
eurt
s (m
ijn o
ud-s
tage
bege
leid
er
goed
e ge
spre
kken
! Be
halv
e on
ze e
igen
kan
toor
kam
er zi
jn e
r bij
onde
rzoe
kcen
trum
B-W
ARE
natu
urlij
k no
g ve
el
uite
rlijk
schu
ilt e
en k
lein
en
heel
zorg
zaam
har
tje. O
ok R
ick
Kuip
erij
kan
hier
nie
t ont
brek
en,
oplo
ssin
gen
zijn
van
onsc
hatb
are
waa
rde.
Uite
raar
d zij
n er
nog
een
hel
eboe
l ana
liste
n m
eer
11
138
erg
beda
nkt v
oor j
ullie
bijd
rage
aan
ond
er a
nder
e IC
P, A
uto
Anal
yzer
, C:N
, am
inoz
uur e
n w
orte
l an
alys
es. M
aar o
ok d
e an
alist
en b
ij B-
WAR
E; D
eef v
an H
oudt
, Kar
en B
onge
rs, J
ohn
Pete
rs, D
olf
hebb
en d
oor d
e ja
ren
heen
vel
e bo
dem
- en
wat
erm
onst
ers v
erw
erkt
! Van
de
afde
ling
Ecol
ogie
de R
adbo
ud U
nive
rsite
it N
ijmeg
en h
eb ik
vee
l gas
-ana
lyse
s mog
en u
itvoe
ren
en h
ierb
ij te
kst e
n
heel
erg
bed
ankt
voo
r jul
lie m
eede
nken
, vel
e hu
lp, l
ekke
re th
ee e
n ge
zelli
ghei
d! Ik
hoo
p no
g
geho
lpen
bij
mijn
ond
erzo
ek d
oor g
oede
stud
ente
n di
e al
len
in m
eer o
f min
dere
mat
e he
bben
ik w
ens j
ullie
alle
maa
l hee
l vee
l suc
ces m
et ju
llie
verd
ere
loop
baan
en
wil
julli
e hi
erbi
j nog
maa
ls be
dank
en v
oor j
ullie
hul
p en
inze
t! Zo
als i
k al
zei h
eb ik
aan
een
aan
tal p
roje
cten
kun
nen
wer
ken
bij B
-WAR
E, w
aaro
nder
het
Zijp
, Mar
cel G
root
en
Andr
é Ti
mm
er v
an N
atuu
rmon
umen
ten,
land
eige
nare
n va
n de
ond
erzo
ek
perc
elen
en
alle
vrij
will
iger
s wil
ik h
ierb
ij be
dank
en v
oor j
ullie
insp
anni
ngen
en
med
ewer
king
. Een
an
der l
angd
urig
pro
ject
was
de
beka
lkin
gspr
oef i
n de
Nie
uwko
opse
pla
ssen
, die
staa
t bes
chre
ven
en m
edew
erki
ng h
ierin
. Voo
r hoo
fdst
uk 3
en
4 he
b ik
pla
nten
mog
en v
erza
mel
en in
de
Bruu
k,
Rijn
land
en h
ier n
og sp
ecia
al b
edan
ken
voor
inzic
hten
die
ik h
eb v
erkr
egen
uit
de le
uke
proj
ecte
n
het o
psch
rijve
n va
n de
ond
erzo
ek re
sulta
ten.
Ook
hie
rbij
heb
ik v
an v
eel p
erso
nen
hulp
mog
en
hier
in! T
ijden
s het
schr
ijfpr
oces
heb
ik o
ok g
oede
feed
back
en
waa
rdev
olle
stof
tot n
aden
ken
gekr
egen
van
Em
iel B
rouw
er e
n Ro
land
Bob
bink
van
Ond
erzo
ekce
ntru
m B
-WAR
E en
van
Eric
riem
wor
dt g
esto
ken,
en
hier
voor
wil i
k ie
dere
en va
n m
ijn co
llega
’s bi
j B-W
ARE
en va
n de
Eco
logi
e-gr
oep
voor
zove
r ik
die
nog
niet
bij n
aam
had
gen
oem
d al
lem
aal b
ijzon
der v
oor b
edan
ken!
Dit
geld
t oo
k bui
ten
de w
erkm
uren
, wan
t alle
lieve
vrie
nden
, fam
ilie e
n ke
nniss
en b
inne
n en
bui
ten
de R
eek
Een
bijzo
nder
mom
ent v
oor o
ns b
eide
n, b
edan
kt v
oor j
ouw
ver
trou
wen
en
bijd
rage
! Leo
n, ji
j
sche
rp e
n na
uw b
etro
kken
, maa
r ook
hee
l soc
iaal
en
mee
leve
nd w
anne
er d
at n
odig
is. I
k de
nk
in h
et la
b op
je h
oofd
, fel
blau
we
eten
swar
en te
sten
, tro
pisc
he b
lous
es, e
n de
kle
ts o
ver o
nze
verd
er li
et g
aan.
Jij b
ent d
egen
e di
e m
ij na
het
afs
tude
ren
aan
deze
leuk
e ba
an b
ij B-
WAR
E he
bt
geho
lpen
, en
ik h
oop
dat j
ouw
ver
trou
wen
er n
og st
eeds
is n
u w
e hi
er st
aan.
Dit
laat
ste
geld
t ook
voor
me
klaa
r sto
nden
, voo
r jul
lie e
ntho
usia
sme
en o
nmisb
are
steu
n. Ik
wee
t nie
t goe
d ho
e te
la
ten
wet
en h
oe d
ankb
aar i
k be
n vo
or a
lles,
dus
daa
rom
laat
ik h
et h
ier b
ij en
ver
trou
w ik
ero
p da
t jul
lie h
et w
el w
eten
. Ik
had
me
geen
bet
ere
bege
leid
ers k
unne
n w
ense
n, e
n ne
em ju
llie
zove
r ov
erge
drag
en k
enni
s mee
naa
r een
hop
elijk
moo
ie to
ekom
st a
ls ‘m
odde
rdoc
tor’
!
139
11
BIOG
EOCH
EMIC
AL W
ATER
-MAN
AGEM
ENT
& AP
PLIE
D RE
SEAR
CH O
N EC
OSYS
TEM
S
Onde
rzoe
kcen
trum
B-W
ARE
Onde
rzoek
centr
um B-
WARE
B.V.
is een
spin-
off be
drijf v
an de
afde
ling A
quati
sche E
colog
ie &
Milie
ubiol
ogie,
welk
e dee
l uitm
aakt
van h
et on
derzo
ekins
tituu
t IWW
R (Ins
titute
for W
ater a
nd W
etlan
d Re
searc
h) va
n de R
adbo
ud U
nivers
iteit N
ijmeg
en. B
-WAR
E werd
in no
vemb
er 20
02 op
geric
ht en
co
mbine
ert de
expe
rtise
van z
uiver
weten
schap
pelijk
en to
egep
ast u
nivers
itair o
nderz
oek.
Kenn
isont
wikk
eling
en -u
itwiss
eling
tusse
n de U
nivers
iteit e
n B-W
ARE i
s het
uitga
ngsp
unt e
n staa
t bo
rg vo
or ho
ogwa
ardig
toege
past
onde
rzoek
, noo
dzak
elijk
voor
het o
plosse
n van
prak
tijkvra
gen
van o
pdrac
htgev
ers.
Spec
ialis
atie
De w
erkwi
jze be
staat
uit de
comb
inatie
van a
nalys
es in
de ve
ldsitu
atie e
n het
opze
tten e
n uitv
oeren
van
velde
xperi
mente
n en e
xperi
mente
n ond
er ge
contr
oleerd
e lab
orator
iumco
nditie
s, naa
st de
skres
earch
. De
verkr
egen
resu
ltaten
drag
en di
rect b
ij aan
het o
ntwi
kkele
n van
nieu
we in
zichte
n binn
en he
t natu
ur-
en w
aterb
ehee
r en w
orden
bijvo
orbee
ld toe
gepa
st bij
natuu
rhers
tel en
–on
twikk
eling
, en w
aterb
erging
. De
ze w
erkwi
jze m
aakt
het m
ogeli
jk da
t de a
dviez
en bi
j uitv
oerin
g in d
e prak
tijk ee
n grot
e kan
s van
sla
gen h
ebbe
n. Er
wordt
onde
rzoek
verric
ht aa
n stur
ende
proc
esse
n in o
nder
meer
hoog
vene
n, laa
gven
en,
meren
, plas
sen,
venn
en, b
eekd
alen,
heide
n, so
orten
rijke g
raslan
den,
polde
rs en
broe
kbos
sen.
Voor
wie
? On
derzo
ekce
ntrum
B-WA
RE ve
rrich
t ond
erzoe
k vo
or mi
nister
ies, p
rovinc
ies, p
rovinc
iale l
ands
chap
pen,
geme
enten
, part
iculie
re ter
reinb
ehee
rders
en na
tuur-
besch
erming
sorga
nisati
es, w
atersc
happ
en
en ad
viesb
ureau
s.
TE
ww
w.b-
war
e.eu
