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.

Isnt

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

180

coun

trie

s (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

vre

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

the

pea

t is

an im

port

ant

& 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

32S

O4

et a

l.,

42-

4+ a

nd N

O3-

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- o

r SO

42-

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- le

vels

may

also

incr

ease

as a

re

sult

of th

e di

scha

rge

of g

roun

dwat

er fr

om su

rrou

ndin

g fo

rest

s or a

gric

ultu

ral a

reas

.

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

+ , S2

- or

2+et

al.,

199

5; L

ucas

sen

et

al.,

2002

; Lam

ers

et a

l4+

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 re

leas

e fr

om 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- a

vaila

bilit

y

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 int

rusio

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

m

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

the

surfa

ce w

ater

. Dur

ing

both

Ch

emic

al a

naly

ses

cont

aini

ng 0

.2 M

H3P

O4

22

dire

ctly

tran

spor

ts th

e CO

2

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-

(Kam

phak

e et

al

4+

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

m

ol L

-1 fo

r se

dim

ent A

, and

bet

wee

n 0-

500 m

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 m

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

bein

g m

obili

sed

from

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 a

nd

2 ra

tes a

nd 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

rea

dily

tak

en u

p as

a n

utrie

nt, s

ever

al

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-

1-1

4+

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+ l

oadi

ngs i

n th

e sh

ort t

erm

,

4+ lo

adin

gs.

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 r

eadi

ly 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+ ri

ch, a

cidi

c sy

stem

s ar

e m

ore

tole

rant

to

high

NH 4

+

4+ (D

e Gr

aaf e

t al

et a

l.,et

al

4+

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+

avai

labi

lity

for

mar

shes

, a

nd B

row

n se

dge

(te

sted

whe

ther

thes

e w

etla

nd sp

ecie

s ind

eed

pref

er th

e up

take

of N

H 4+ r

athe

r tha

n N

O3-

. Sec

ondl

y, th

eir

resp

onse

s an

d to

lera

nces

to

stro

ngly

inc

reas

ed N

H 4+

4+ to

lera

nce

of w

etla

nd

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

1o46

N; 5

o

into

an

aera

ted

18

3- o

r NH 4

+

adju

stm

ent o

f 7 d

ays

in a

erat

ed 1

8 bo

th N

O3-

and

NH 4

+10

0 m

ol L

-1 C

a2+ ,

100 m

ol L-

1 Mg

2+, 2

00

mol

L-1 K

+ , 10

0 m

ol L-

1 SO

42- , 2

00

mol

L-1 P

O43

- , 0.8

m

ol L-

1 Mn2

+ , 0.

8 m

ol

L-1 H

3BO

3, 0.

7 m

ol L

-1 Z

n2+ ,

0.27

m

ol L

-1-1

Cu2

+ , 0.

008 m

ol L

-1

Mo

(De

Graa

f et a

l.,, w

ith b

oth

NO

3- (2

0 m

ol L

-14+

(20 m

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 . Th

ree

indi

vidu

als o

f the

sam

e sp

ecie

s wer

e pl

aced

in e

ach

plan

t con

tain

er b

y us

ing

eith

er N

a15 N

O3 o

r 15

NH 4

Cl (

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 m

ol L

-1 N

H 4+ a

nd 2

mm

ol L

-1 N

H 4+

usin

g 1.

2 m

ol L-

1 HCl

and

1 m

ol L-

1Th

e -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 15 N

.

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+ v

ersu

s NO

3Bo

th

and

w

ere

able

to ta

ke u

p N

H 4+ as

wel

l as

NO

3- at

bot

h pH

4 a

nd p

H 6 4+

than

for N

O3-

4+3-

in

and

4+

spec

ies,

incr

ease

d N

H 4+

-l N

H 4+

-1

NH4+

betw

een

pH a

nd h

igh

NH 4

+ was

foun

d in

sho

ot b

iom

ass.

This

led

to a

n in

crea

se o

f sho

ot b

iom

ass

in

the

high

NH 4

+ tr

eatm

ent c

ompa

red

to th

e lo

w N

H 4+

4+

4+ a

nd p

H w

as

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+

trea

tmen

t in

shoo

t

tre

ated

with

hi

gh N

H 4+ .

In ro

ot b

iom

ass

of 4+

and

high

pH.

s

how

ed

42

L4

4, a

t

shoo

t bio

mas

s, rh

izom

e bi

omas

s

43

mol L4mol L4

3

4+ a

nd p

H, a

nd a

n in

crea

se in

shoo

ts o

nly

with

hig

h N

H 4+

, pH

and

NH 4

+

4+ a

nd lo

w p

H.

4+

4+ a

nd

low

pH.

4+ a

t low

pH,

rega

rdle

ss o

f the

N tr

eatm

ent.

Mag

nesiu

m

4+ a

nd a

t a

4+ tr

eatm

ents

wer

e fo

und.

an

incr

ease

of s

hoot

bio

mas

s at

hig

h N

H 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+

trea

tmen

t. Th

e do

min

ant

N-r

ich

free

am

ino

acid

was

asp

arag

ine,

whi

ch a

ccou

nted

alre

ady

for 7

1-73

% o

f the

tota

l fre

e am

ino

. Alth

ough

show

ed a

sim

ilar

resp

onse

to

enha

nced

NH 4

+. J

.

incr

ease

d le

ss th

an in

4+ a

nd p

H. H

owev

er,

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

an

ener

gy d

eman

ding

pro

cess

for p

lant

s, b

ecau

se it

has

to b

e re

duce

d to

NH 4

+ Be

caus

e N

H 4+

4+ r

ich

envi

ronm

ents

, a

pref

eren

ce o

f N

H 4+

, and

to

a 4+

. Alth

ough

NH 4

+ upt

ake

was

hig

her,

NO

3-

was

also

tak

en u

p re

adily

, esp

ecia

lly b

y .

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- o

r NH 4

+ (Ch

ang

et a

l3-

, also

has

the

high

est r

adia

l et

al

3-4+

are

also

4+ t

reat

men

t (2

mm

ol

L-1Ac

orus

cal

amus

et a

l.an

d (T

ylov

a et

al.,

4+ lo

adin

gs

(res

p. 1

4.8

& 3

.7 m

mol

L-1

and

in

4+

. Thi

s abi

lity

to c

hang

e on

ly b

iom

ass

4+ t

oler

ance

. Suc

h a

tole

ranc

e to

hi

gh N

H 4+ l

oadi

ngs i

n w

etla

nd g

ram

inoi

ds w

as so

far o

nly

foun

d fo

r Phr

agm

ites a

ustr

alis

(Tyl

ov

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+ t

reat

men

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

ov

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+ l

evel

s.

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+ ,

beca

use

it is

acco

mpa

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+

at p

H 4,

but

not

at p

H 6,

due

to in

tern

al

4+ u

ptak

e ra

tes

of b

oth

test

ed g

ram

inoi

ds w

ere

4+ is

mos

t

46

incr

easin

g N

H 4+

, Ca

lluna

vul

garis

, (V

an d

en B

erg

et a

l.,in

Rici

nus

com

mun

is (V

an B

eusic

hem

et a

l.,

(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+. O

n th

e ot

her h

and,

suc

h ch

ange

s in

pla

nt m

orph

olog

y an

d ph

ysio

logy

as a

resu

lt of

incr

ease

d N

H 4+

et a

l A

lthou

gh a

redu

ced

root

:sho

ot

NH 4

+ et

al.,

, fo

und

for b

oth

gram

inoi

ds in

resp

onse

to e

nhan

ced

NH 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+ d

irect

ly

(Van

Dijk

& R

oelo

fs, 1

988;

Rab

e, 1

990;

Ns

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+ l

evel

s. 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+ u

ptak

e, w

hile

gr

owth

was

not

enh

ance

d, h

owev

er,

acid

N c

ompo

unds

that

wer

e no

t mea

sure

d in

our

stud

y. C

ruz

et a

l.pl

ant t

oler

ance

to N

H 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

(or

even

dee

per

for

tree

s; C

anad

ell e

t al

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,

NO3-

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

supp

lied

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-

and

OH-

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+ o

r N

O3-

, 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+

upt

ake,

alth

ough

NH 4

+

3-. P

lant

s of r

ipar

ian

wet

land

s, h

owev

er, a

re k

now

n to

face

pro

blem

s at

a hi

gh N

H 4+ :

NO

3-et

al

., 20

03; V

an d

en B

erg

et a

l4+

inpu

ts, a

s a re

sult

of a

mm

onia

(N

H 3et

al

4+ :

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 e

t al.,

199

8; B

urgi

n an

d Ha

milt

on, 2

007;

Hu

et a

lTh

e up

take

of N

H 4+ o

r NO

3-

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

ord

er to

cha

nge

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- an

d S2

- , pK

12

H 2S

will

be

prev

ailin

g du

e to

HCO

3CO

2

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- (H

aaije

r et

al

orga

nic

com

poun

ds, N

O3-

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 S

phag

num

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 (

aver

age

size

of 1

0 m

2

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

lev

, div

ided

by

the

log

of th

e st

anda

rd re

lev

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

H3P

O4

2

22

2 and

HCO

3-

ml 6

5% H

NO

3-

2+, M

g2+

3- (K

amph

ake

et a

l4+

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

&

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

vs 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

vs

gene

rally

onl

y co

ntai

ned

com

mon

spe

cies

.

S. p

alus

tre

S.

,

. In

only

3 re

lev

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+

avai

labl

e N

O3-

plan

t av

aila

ble

K+

43- ,

NO3-

, NH 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

vs,

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

evs

; 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.,

lso

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 (

Ml

son

et a

l., 2

010;

Hed

berg

et a

l., 2

012;

Mid

dlet

on e

t al.,

et

al.,

4+

et

alet

al

of N

H 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- t

o ad

jace

nt s

urfa

ce w

ater

s,

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