Wetland Restoration With Plants

7/23/2019 Wetland Restoration With Plants

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WETLANDS,

Vol. 17, No. 3 , September 1997, pp. 424-437

© 1997 , The Soc ie ty o f Wet land Sc ien t i s t s

R E S T O R A T I O N O F W E T L A N D V E G E T A T I O N W I T H T R A N S P L A N T E D

W E T L A N D S O I L : A N E X P E R I M E N T A L S T U D Y

S t e p h e n C . B r o w n ~ a n d B a r b a r a L . B e d f o r d

Department of Natural Resources

Cornell University, Fernow Hall

Ithaca, NY 14853

1Present address:

Department of Plant and Soil Science

University of Massachusetts, Stockbridge Hall

Amherst, MA 01003

e-mail Stephen. Brown @ state.ma.us.

Abstract:

R e s t o r a ti o n o f d r a i n e d w e t l a n d s r e q u i r es t h e r e - e s t a b l i s h m e n t o f a n a t i v e w e t l a n d p l a n t c o m m u -

n i t y . T h i s c a n b e d i f f i c u l t i n a re a s w h e r e l o n g - t e r m d r a i n a g e h a s e l i m i n a t e d w e t l a n d v e g e t a t i o n a n d s i g n i f i -

c a n t l y r e d u c e d t h e n u m b e r o f v i a b l e w e t l a n d p l a n t s e e d s i n t h e s e e d b a n k . T h i s s t u d y o f U . S . F i s h a n d

W i l d l i f e S e r v i c e w e t l a n d r e s t o ra t i o n s i t e s in n o r t h e r n N e w Y o r k e x a m i n e s t h e e f f e c t i v e n e s s o f t r a n s p la n t i n g

w e t l a n d s o i l f r o m s m a l l r e m n a n t w e t l a n d s i n t h e d r a i n a g e d i t c h es t o t h e a r e a th a t b e c o m e s s h a l l o w m a r s h

f o l l o w i n g r e f l o o d i n g . T h e r e s u l t s o f t w o e x p e r i m e n t s a r e r e p o r t e d , i n c l u d i n g a s m a l l - s c a l e s t u d y o f t r a n s -

p l a n t a t i o n t e c h n i q u e s u s i n g s m a l l p l o t s w i t h t r e a t m e n t s a n d c o n t r o l s e s t a b l i s h e d b y h a n d , a n d a l a r g e - s c a l e

a p p l i c a t i o n o f s o i l t r a n s p l a n t a t i o n a n d s i t e - p r e p a r a t i o n te c h n i q u e s u s i n g h e a v y e q u i p m e n t t o e s t a b l i s h l a r g e

p l o t s a c r o s s e n t i r e w e t l a n d b a s i n s . I n t h e s m a l l - s c a l e s t u d y , t h e t r a n s p l a n t p l o t s h a d s i g n i f i c a n t l y l o w e r

w e t l a n d i n d e x v a l u e s , i n d i c a t i n g g r e a t e r d o m i n a n c e o f w e t l a n d p l a n t s , a f te r o n e g r o w i n g s e a s o n b u t n o t a f te r

t w o . T r a n s p l a n t p lo t s a l s o h a d m o r e w e t l a n d p l a n t s p e c i e s a n d m o r e w e t l a n d p l a n t c o v e r t h a n n a t u r a l c o n t ro l

p l o t s , a n d t h e s e d i f f e r e n c e s p e r s i s t e d t h r o u g h t h e s e c o n d g r o w i n g s e a s o n . L i t t e r r e m o v a l a n d s o i l d i s t u r b a n c e

a l s o l o w e r e d t h e w e t l a n d i n d e x v a l u e s a n d i n c r e a s e d w e t l a n d p l a n t s p e c i e s n u m b e r a n d c o v e r , b u t o n l y f o r

t h e f i rs t g r o w i n g s e a s o n . I n t h e l a r g e - s c a l e s t u d y , s o i l t r a n s p l a n t a t i o n s i g n i f i c a n t l y i n c r e a s e d b o t h t h e n u m b e r

o f s p e c i e s a n d t h e a m o u n t o f c o v e r o f w e t l a n d p l a n t s a n d o f p l a n t s v a lu a b l e a s w i l d l i fe f o o d s o u r c e s . M o w i n g

a n d p l o w i n g t r e a t m e n t s i n c r e a s e d w e t l a n d p l a n t e s t a b l i s h m e n t , b u t m u c h l e s s t h a n s o i l t r a n s p l a n t a t i o n , a n d

p l o w i n g s i g n i f i c a n t l y i n c r e a s e d t h e e s t a b l i s h m e n t o f c a t t a i l

Typha

s p p . ) , a n u n d e s i r a b l e i n v a s i v e s p e c i e s i n

s m a l l w i l d l i f e m a r s h e s . S o i l t r a n s p l a n t a t i o n s h o u l d b e a p a r t i c u l a r l y e f f e c t i v e t e c h n i q u e f o r i m p r o v i n g w e t l a n d

p l a n t e s t a b l i s h m e n t a n d l i m i t i n g c a t t a i l e n c r o a c h m e n t i n a r e a s d i s t u r b e d b y d i k e c o n s t r u c t i o n ,

Key Words:

w e t l a n d r e s to r a t i o n , w e t l a n d p l a n t s , s o i l a m e n d m e n t s , s o i l t ra n s p l a n t a t i o n , N e w Y o r k

I N T R O D U C T I O N

T h r o u g h o u t t h e U n i te d S t a t e s, g o v e r n m e n t a g e n c i e s

r e q u i r e th a t w e t l a n d s d e s t r o y e d b y v a r i o u s h u m a n a c -

t i v i ti e s b e r e p l a c e d w i t h c r e a t e d o r re s t o r e d s i t es . H o w -

e v er , f e w s t u d i e s h a v e s y s t e m a t i c a l l y e x a m i n e d t h e e f-

f e c t i v e n e s s o f d i f f e r e n t t e c h n i q u e s u s e d t o r e c r e a t e n a t-

u r a l w e t l a n d s . W e f o u n d i n a n o n g o i n g s t u d y o f w e t -

l a n d s r e s t o r e d b y t h e U . S . F i s h a n d W i l d l i f e S e r v i c e

( B r o w n 1 9 9 5 ) t h a t c a t t a i l Typha s p p .) o f t e n d o m i n a t e s

a r e a s o f t h e s i t e s w h e r e c o n s t r u c t i o n a c t i v i ti e s d i s tu r b

t h e s o i l d u r i n g r e s t o r a t i o n . O d u m ( 1 9 88 ) , R e i n a r t z a n d

W a r n e ( 1 9 9 3 ) , a n d a r e v i e w o f w e t l a n d r e s t o r a t i o n b y

t h e N a t i o n a l R e s e a r c h C o u n c i l ( 1 9 9 2) a l s o r e p o r t t h e

t e n d e n c y o f m o n o t y p i c s t a n d s o f c a t t ai l a n d o t h e r a g -

g r e s s i v e n a t i v e a n d e x o t i c s p e c i e s t o p r e c l u d e e s t a b -

l i s h m e n t o f d i v e r s e p l a n t c o m m u n i t i e s i n r e s t o r e d w e t -

l a n d s .

C a t t a i l i n v a s i o n i s p r o b l e m a t i c b e c a u s e d e n s e m o n o -

c u l t u r e s r e d u c e t h e r a ti o o f o p e n w a t e r t o v e g e t a t i o n ,

r e d u c e t h e d i v e r s i t y o f v e g e t a t i o n , a n d t h e r e f o r e r e d u c e

t h e h a b i t a t v a lu e o f t h e s it e ( W e l l e r 1 9 7 5 , K a m i n s k i

a n d P r i n c e 1 9 8 1 , M u r k i n e t al . 1 9 8 2 ). I n a d d i t i o n , c a t -

t a i l s a r e v e r y d i f f i c u l t t o c o n t r o l o n c e t h e y a r e e s t a b -

l i s h e d , e v e n w i t h i n t e n s i v e m a n a g e m e n t t e c h n i q u e s

l i k e c u tt i n g , b u r n in g , a n d t h e u s e o f h e r b i c i d e s ( B e u l e

1 9 79 , M a l l i k a n d W e i n 1 9 8 6 , C o m e s a n d K e l l y 1 9 8 9,

B a l l 1 9 9 0 ) .

G o v e r n m e n t a l a g e n c ie s i n v o l v e d i n re s t o r a ti o n

sen-

s u L e w i s 1 9 8 9) o f w e t l a n d s n e e d r e s t o r a t i o n t e c h -

n i q u e s t h a t i n c r e a s e t h e l i k e l i h o o d o f e s t a b l i s h i n g d i -

4 2 4



B r o w n & B e d f o r d , W E T L A N D R E S T O R A T I O N W I T H T R A N S P L A N T E D S O I L 4 25

v e r s e p l an t c o m m u n i t i e s t h a t r e s e m b l e n a t u r a l w e t -

l a n d s ( N a t i o n a l R e s e a r c h C o u n c i l 1 9 9 2 ). I f i n c r e a s i n g

a v i f a u n a h a b i t a t i s a m o n g t h e r e s t o r a t i o n g o a l s , t h e

t e c h n i q u e s m u s t e s t a b l is h p l a n t c o m m u n i t i e s t h a t e n -

h a n c e h a b i t a t v a l u e f o r b ir d s . F u r t h e r m o r e , t h e te c h -

n i q u e s m u s t h a v e t h e c h a r a c t e r is t i c t h a t t h e y c a n b e

a p p l i e d t o m a n y w e t l a n d s a t th e s c a l e o f e n t ir e re s t o -

r a t i o n s it e s a n d w i t h o u t e x c e s s i v e c o s t t o t h e a g e n c y

o r l a n d o w n e r .

N u m e r o u s s t u d ie s o f n a tu r a l w e t l a n d s h a v e t i e d th e

m a i n t e n a n c e o f a d i v e r s e f l o ra t o th e p r e s e n c e o f a d i -

v e r s e s e e d b a n k t h a t r e s p o n d s t o c h a n g i n g w a t e r l e v e ls

( v a n d e r V a l k a n d D a v i s 1 9 7 8, v a n d e r V a l k 1 9 8 1, P e d -

e r s o n a n d v a n d e r V a l k 1 9 8 4, K e d d y a n d E l l i s 1 9 8 5 ).

S e e d b a n k s w i l l n o t c o n t r ib u t e t o r e s t o r a ti o n o f w e t l a n d

p l a n t c o m m u n i t i e s , h o w e v e r , i f th e y h a v e b e e n s i g n if i -

c a n t l y d e p l e t e d b y l o n g - t e r m d i s t u r b a n c e o f th e f o r m e r

w e t l a n d s it e. A f t e r m a n y y e a r s o f d r a in a g e , f o r e x a m p l e ,

t h e s e e d b a n k i n a f o r m e r w e t l a n d c a n b e c o m e r e la -

t i v e l y d e p a u p e r a t e o f w e t l a n d s p e c i e s ( W e i n h o l d a n d

v a n d e r V a l k 1 9 8 9 , B r o w n 1 9 9 5, G a l a t o w i t s c h a n d v a n

d e r V a l k 1 9 9 6 ) . T h i s l o s s o f v ia b l e s e e d s m a y s l o w o r

p r e v e n t r e c o v e r y o f w e t l a n d v e g e t a t i o n f o l l o w i n g r e s -

t o r a ti o n o f w e t l a n d w a t e r l e v e l s. D e s i r a b l e s p e c i e s m a y

n o t r e - e s ta b l i sh b e f o r e a m o r e a g g r e s s i v e s p e c i e s p r e -

e m p t s t h e s it e ( R e i n a r t z a n d W a r n e 1 9 9 3 ) .

O n s i te s d r a i n e d w i t h d i tc h e s , h o w e v e r , a r e m n a n t

w e t l a n d p l a n t c o m m u n i t y o f t e n e x i s t s i n t h e p o o r l y

d r a i n e d s o i l a d j a c e n t t o t h e d i t c h . W e h y p o t h e s i z e d

t h a t, i f th e s e r e m n a n t s o i ls c o u l d b e t r a n s p l a n t e d t o t h e

i n t e n d e d r e s t o r a t i o n a r e a , th e i r r e l a t iv e l y r i c h s e e d

b a n k s w o u l d p r o d u c e g r e a t e r s p e c i e s r i c h n e s s a n d c o v -

e r o f w e t l a n d p l a n t s f o l l o w i n g r e s t o r a t i o n o f w e t l a n d

h y d r o l o g y t h a n t h e s e e d b a n k s o f a r e as t h a t h a d b e e n

e f f e c t i v e l y d r a i n e d ( D u n n a n d B e s t 1 9 8 4 ) . W e f u r t h e r

h y p o t h e s i z e d t h a t th e t e c h n iq u e o f m o v i n g r e m n a n t

w e t l a n d s o i ls w a s o n e t h a t c o u ld b e i m p l e m e n t e d o n

a s c a l e l a r g e e n o u g h t o a f f e c t e n t i r e r e s t o r a t i o n s i te s .

O u r g o a l w a s t o d e t e r m i n e i f t r a n sp l a n ti n g r e m n a n t

w e t l a n d s o i l c o u l d b e a n e f f e c t i v e t e c h n i q u e f o r r e c -

r e a t i n g a w e t l a n d p l a n t c o m m u n i t y a t h i g h e r e l e v a t i o n s

w h e r e d r a i n a g e w a s m o r e c o m p l e t e . W e c o n d u c t e d o u r

e x p e r i m e n t s a t t w o s c a l e s , a s m a l l - s c a l e s t u d y u s i n g

p l o t s w i t h t r e a t m e n t s e s t a b l i s h e d b y h a n d , a n d a l a r g e -

s c a l e s tu d y u s i n g c o n s t r u c t i o n e q u i p m e n t t o e s t a b li s h

t r e a t m e n t a r e a s a c r o s s e n t i r e w e t l a n d b a s i n s . B e c a u s e

t h e g o a l o f t h e re s t o r a t i o n p r o g r a m w e s t u d i e d w a s t o

i n c r e a s e a v i f a u n a h a b i t a t , w e w e r e p a r t i c u l a r ly i n t e r-

e s t e d i n t h e r e s p o n s e s o f w e t l a n d p l a n t s k n o w n t o b e

u s e d a s f o o d s o u r c e s b y w a t e r f o w l a n d m a r s h b i r d s .

O u r e x p e r i m e n t s w e r e d e s i g n e d t o a d d r e s s t h e fo l l o w -

i n g m a j o r q u e s t i o n s : 1 ) c a n t h e n u m b e r o f w e t l a n d

p l a n t s p e c i e s t h a t g e r m i n a t e a n d e s t a b l i s h b e i n c r e a s e d

b y t r a n s p l a n t i n g s o i l f r o m r e m n a n t w e t l a n d a r e a s ; 2)

d o e s t h e i n c l u s i o n o f w e t l a n d p l a n t r h i z o m e s i n th e

t r a n s p l a n t e d s o i l a f f e c t t h e s p e c i e s r i c h n e s s a n d a b u n -

d a n c e o f w e t l a n d p l a n ts t h a t b e c o m e e s t a b l is h e d ; 3 )

h o w l o n g d o t r e a t m e n t e f f e c t s p e r s i s t ; 4 ) d o t h e t r e a t -

m e n t s i n c r e a s e th e a b u n d a n c e o f p l a n t s p ec i e s k n o w n

t o b e u s e d a s f o o d s o u r c e s b y w a t e r f o w l a n d m a r s h -

b i r d s ; a n d 5 ) c a n t h e s e t e c h n i q u e s b e a p p l i e d a t t h e

s c a l e o f w h o l e r e s t o r a t i o n s i t e s ?

M E T H O D S

S m a l l - S c a l e S t u d y

W e c o n d u c t e d t h e s m a l l - s c a l e s t u d y o n s i x re s t o r a -

t i o n s i te s s e l e c t e d f r o m a t o t a l o f 2 1 i n o u r o n g o i n g

s t u d y o f v e g e t a ti o n r e s p o n s e s t o re s t o ra t i o n ( B r o w n

1 9 9 5 ) . T h e s t u d y s i te s a r e l o c a t e d i n J e f f e r s o n C o u n t y ,

N e w Y o r k ( 4 4 ° 0 0 'N , 7 5 ° 5 2 ' W ) i n th e L a k e O n t a r io

a n d S t. L a w r e n c e R i v e r l o w l a n d s . A t e a c h s t u d y s i te ,

G u f f i n s e r i e s s o i l s o c c u p y t h e l o w e s t a r e a s ( a f in e ,

m i x e d , n o n a c i d , m e s i c M o l l i c E p i a q u e p t ) g r a d i n g t o

C h a u m o n t s e r i e s s o il s i n t h e s u r r o u n d i n g u p l a n d s ( a

f in e , m i x e d , m e s i c A e r i c E p i a q u a l f ) .

E a c h r e s t o r a t i o n s i te h a d a r e m n a n t w e t l a n d c o r r i d o r

l o c a t e d i n t h e d r a i n a g e d i t c h c o n s t r u c t e d t o r e m o v e

w a t e r f r o m t h e b a si n . D r a i n a g e o c c u r r e d a t l e a s t 4 0

y e a r s a g o a t al l s i t es a n d w a s p r o b a b l y c o m p l e t e d

m u c h l o n g e r a g o a t m o s t s i t e s, b u t n o r e c o r d s a r e a v a i l-

a b le . T h e r e m n a n t w e t l a n d c o r r i d o r s h a v e m a n y m o r e

w e t l a n d p l a n t s p e c i e s a n d s p e c i e s - r i c h s e e d b a n k s t h a n

t h e re m a i n i n g d r a i n e d a r e a s t h a t b e c o m e s h a l l o w

m a r s h f o l l o w i n g r e s t o r a ti o n o f w e t l a n d h y d r o l o g y

( B r o w n 1 9 9 5 ). T h e o r i g i n a l w e t l a n d v e g e t a t i o n a t

h i g h e r e l e v a t i o n s w a s e n t i r e l y r e p l a c e d b y u p l a n d f i e ld

p l a n t s , as d r a i n e d a r e a s b e c a m e t o o d r y t o s u p p o r t h y -

d r o p h y t i c v e g e t a t i o n . T h e d r a i n e d a r e a s w e r e u s e d p r i-

m a r i l y f o r p a s t u r e a n d f o r a g e c r o p s , w i t h v e r y l i tt l e

c u l t i v a t i o n .

T h e e x p e r i m e n t c o n s i s t e d o f t w o t r e a t m e n t s a n d

t h r e e t y p e s o f c o n t r o l s , a ll o f w h i c h w e e s t a b l i s h e d i n

M a y o f 1 9 9 2 ( T a b l e 1 A ) . A t e a c h o f th e s i x r e s t o r a t i o n

s i te s , p l o t s w e r e e s t a b l i s h e d a t 1 5 c m b e l o w t h e r e -

s t o re d m a x i m u m w a t e r l ev e l b y s u r v e y i n g w i t h a n

a u t o - l e v e l . I n t h e f ir s t t r e a t m e n t , w e t r a n s p l a n t e d s o i l

b l o c k s b y r e m o v i n g a l l u p l a n d s o i l in 0 . 2 5 m 2 p l o t s t o

a d e p t h o f 1 5 c m , t h e n t r a n s p l a n t i n g a l l s o i l, p l a n t

r o o t s , a n d r h i z o m e s f r o m a n e q u a l s o i l v o l u m e in t h e

r e m n a n t w e t l a n d t o th e e x p e r i m e n t a l p l o t. W e t ra n s -

p l a n t e d t h e s o i l i n f o u r a p p r o x i m a t e l y e q u a l s e c t i o n s ,

w h i c h w e r e m o v e d a s i n t ac t a s p o s s ib l e . W e r e p l ic a t e d

t h is t r e a t m e n t t h re e t i m e s a t e a c h o f th e 6 w e t l a n d s . I n

t h e s e c o n d t r e a t m e n t , w e f ir s t s i e v e d t h e t r a n s p l a n t e d

s o il b y p a s s i n g i t th r o u g h a 1 - c m s c r e e n to r e m o v e

p l a n t ro o t s a n d r h i z o m e s . W e t h e n t r a n s p l a n t e d t h e

s i e v e d s o i l i n t h e s a m e w a y a s t h e s o i l b l o c k s . W e

r e p l i c a t e d t h is t r e a t m e n t t h r e e t i m e s a t e a c h o f t w o



426 WE TLA ND S, Volu me 17, No. 3, 1997

Table 1. A) The numbers of quadrats in each treatment type in the small-scale study. (Trans. Blocks = transplanted soil

blocks; Sieved Soil = transplanted sieved soil; Unmanipulated = unmanipulated control plots; Litter Removed = litter removal

without soil disturbance; Disturbed = upland soil removed and replaced. Drawdown plots include the -15 cm plots from 5

of the 6 sites where a natural d rawdown exposed the soil surface.) B) Total numbers of quadrats established in each treatment

type in the large-scale study.

A) Small-Scale Experiment

Plot Type Trans. Blocks Sieved Soil Unmanipulated Litter Remo ved Disturbed

Plots at - 15 cm 18 6 18 6 6

Plots at - 30 cm 0 6 2 2

Total Plots 24 6 24 8 8

Drawdown Plots 15 6 15 6 6

B) Large-Scale Experiment

Treatment Control Mowe d Plowed Soil Trans. Total

Total Plots 39 30 18 54 141

sites. Unmanipulated control plots were established at

the same elevation as the treatment plots, but were not

disturbed in any way, and were replicated three times

at each of the six sites. Litter-removal plots had all

surface litter and plant stems removed. Soil distur-

bance plots had the upland soil to a depth of 15 cm

removed in the same manner as the transplanted soil

blocks, followed by replacement of the same upland

soil. Litter-removal and soil-disturbance plots were es-

tablished once at each of the six sites. To address the

influence of hydroperiod on plant responses to these

treatments, we establis hed an additional set of plots at

30 cm below maximum water level in two sites, each

with three replicates of transplanted soil blocks and

unmanipulated control plots, and one replicate each of

litter removal and soil disturbance plots.

This design made it possible to compare the effects

of the two treatment types and to determine if the ef-

fects resulted from soil disturbance, litter removal, or

from the transplantation of wetland soil. We analyzed

the effect of disturbing the soil by comparing unma-

nipulated controls with disturbed soil plots; the effect

of litter removal by comparing litter removal plots to

unmanipulated controls; the effect of transplanting

wetland soil by comp aring the soil-transplant treatment

with unmanipu lated controls; and the effect of includ-

ing rhizomes in the cores by comparing the sieved

transplanted soil treatment to the transplanted soil

block treatment.

In October of 1992, after seed germination and seed-

ling establishment, and again in September of 1993, we

determined species presence and estimated percent cover

in each plot using a plot frame with measured sides.

Some seedlings were only identifiable to genus, but most

were identifiable to species. We marked unidentified pe-

rermial plants in the field and identified them when flow-

ers or fruits developed. Taxo nomic nomenclature follows

Gleason and Cronquist (1991).

We analyzed the data for each plot by calculating a

weighted average index value (Wentworth and John-

son 1986) based on the wetland indicator status of

each species (Reed 1988) and its importance value in

the plot as determined by its relative percent cover.

Wetland indicator status ranges on a scale from obli-

gate wetland to obligate upland. The categories reflect

the estimated range of probabilities of a species oc-

curring in wetland areas as defined by Cowardin et al.

(1979) as follows: Obligate Wetland (OBL) = >9 9%;

Facultative Wetland (FACW) = 67-99%; Facultative

(FAC) = 34- 66 %; Facultative Upland (FACU) = 1-

33%; Obligate Upland (UPL) = <1%. Many studies

use an index that assigns each class an integer value

from 1 for obligate wetland to 5 for obligate upland

(Siegelquist et al. 1990). Michener (1983) suggested

that an appropriate index should reflect the fact that

the probability classes in the indicator status list are

not evenly spaced between 1 and 100%. We used the

frequency midpoint index value proposed by Eicher

(1988) for calculations on wetland indicator status,

which assigns a value to each class designed to reflect

the midpoint of its probability range, as follows: OBL

= 1.0; FA CW = 1.67; FAC = 3.0; FAC U = 4.33;

UP L = 5.0. Species with an indicat or status of FAC,

FACW, or OBL are considered wetland species, so

lower weighted average index values mean greater

dominance by wetland plants.

We computed a wetland index from the weighted

average indicator values for each plot and compared

plot treatments by analyzing differences in the index

values. We determined the wetland index (WI) of each

vegetation plot according to the formula:

•

( IVl 'WIS1)

i=]

W I =

100



B r o w n & B e d f o r d , W E T L A N D R E S T O R A T I O N W I T H T R A N S P L A N T E D S O I L 4 2 7

T a b l e 2 . W e t l a n d p l a n t s p e c i e s w i t h h i g h v a l u e a s w i l d l i f e f o o d s o u r c e s t h a t o c c u r r e d a t s i t e s in t h i s s t u d y , e = e x c e l l e n t , g

= g o o d , f = f a ir , p = p o o r , b u t i m p o r t a n t f o r s o m e s p e c i e s . C o m p i l e d f r o m M a r t i n e t a l. ( 1 9 5 1 ) a n d P a y n e ( 1 9 9 2 ) .

M a r s h b i r d s a n d

G e n u s S p e c i e s P a r t C o n s u m e d W a t e r f o w l S h o r e b i r d s

A l i s m a s p s e e d s f

B i d e n s s p s e e d s p

C a r e x s p s e e d s f

C o m u s s p f r u i t s g

E l e o c h a r i s s p r o o t s p f

G l y c e r i a s t r i a t a L a m . s e e d s f

J u n c u s s p s e e d s f

L e e r s i a o r y z o i d e s L . s e e d s , r o o t s g

L e m n a m i n o r L . f o l i a g e g

L u d w i g i a p a l u s t r i s L . s e e d s f

P o l y g o n u m h y d r o p i p e r

L . s e e d s g f

P o l y g o n u m s p s e e d s e f

P o t a m o g e t o n s p s e e d s , r o o t s , s t e m s g e

S a g i t t a r i a l a t i f o l i a W i l l d . s e e d s , r o o t s f

S p a r g a n i u m e u r y c a r p u m E n g e l m . s e e d s f f

w h e r e I V = t h e I m p o r t a n c e V a l u e o f s p e c i e s i i n t h a t

p l o t , a n d W I S --- t h e W e t l a n d I n d i c a t o r S t a t u s o f t ha t

s p e c i e s . T h e r e l a t i v e p e r c e n t c o v e r o f e a c h s p e c i e s i n

e a c h p l o t w a s u s e d t o c a l c u l a t e i t s i m p o r t a n c e v a l u e i n

t h a t p l o t. W e s u m m e d t h e p e r c e n t c o v e r o f a l l s p e c i e s

r e c o r d e d i n t h e p l o t a n d d i v i d e d t h e p e r c e n t c o v e r o f

e a c h s p e c i e s b y t h e t ot a l s o i m p o r t a n c e v a l u e s f o r e a c h

p l o t s u m t o 1 0 0 . W e a l s o c o m p a r e d p l o t t y p e s w i t h

r e g a r d t o t h e m e a n n u m b e r o f w e t l a n d s p e c i e s w i t h a n

i n d i c a t o r s t at u s o f F A C , F A C W , o r O B L a n d t h e to t a l

c o v e r o f t h e se w e t l a n d p l a n t s .

W e c o m p a r e d t h e t ra n s p l a n te d w e t l a n d s o i l b l o c k s

a n d u n d i s t u r b e d c on t r o ls w i t h a t w o - w a y A N O V A o n

w e t l a n d i n d e x v a l u e s , w i t h f a c t o r s o f t r e a tm e n t a n d s i t e

( W e t l a n d I n d e x f o r C o v e r = W I ) . W e a l s o t e s t e d f o r

d i f f e r e n c e s i n t h e n u m b e r o f w e t l a n d s p e c i e s p r e s e n t

( W e t l a n d S p e c i e s N u m b e r = W S N ) a n d th e t o t a l p e r -

c e n t c o v e r o f w e t l a n d s p e c i e s i n e a c h p l o t ( W e t l a n d

S p e c i e s C o v e r = W S C ) . E a c h t e st h a d t h r e e r e p l i c a t e

p l o t s p e r t r e a t m e n t i n e a c h o f t h e s i x s i te s a n d t h e t w o

f a c t o r s o f p l o t t y p e a n d s i t e. W e u s e d n o r m a l p r o b a -

b i l i t y p l o t s to t e s t a s s u m p t i o n s o f n o r m a l i t y f o r r e s i d -

u a l s .

A l l o t he r c o m p a r i s o n s w e r e t e s te d u s i ng K r u s k a l -

W a l l i s o n e - w a y a n a l y s e s o f v a r i a n c e b e c a u s e o f t h e

s a m p l e s i z e s in e a c h t r e a t m e n t a n d n o n - n o r m a l i t y o f

d a t a ( W i l k i n s o n 1 9 9 2) . W e r e p e a t e d t h e t es t s f or a l l

t h re e o f t h e r e sp o n s e v a r i a b l e s d e s c r i b e d a b o v e ( W I ,

W S N , a n d W S C ) .

L a r g e - S c a l e S t u d y

W e s e l e c t e d f iv e a d d i t i o n a l d r a i n e d w e t l a n d s i n t h e

s u m m e r o f 1 9 9 3 t ha t w e r e s c h e d u l e d f o r r e s t o r a t i o n

w o r k t h a t f al l . E a c h s i t e h a d s o i l s i n t h e s a m e f a m i l y

a s t h e s i te s i n th e s m a l l s c a l e s t u d y . A t e a c h s i te , w e

e s t a b l i s h e d s o i l t r a n s p l a n t p l o t s b y s u r v e y i n g t h e e l e -

v a t i o n s o f t h e s i te a n d m a r k i n g a n a r e a a t l e a s t 1 0 - m

w i d e a n d e x t e n d i n g a c r o s s t h e e n t i r e w e t l a n d b a s i n

p e r p e n d i c u l a r t o t h e d r a i n a g e c h a n n e l ( b e t w e e n 5 5 a n d

9 5 m a c r o s s ) . W e m a r k e d s i m i l a r l y s i z e d p l o ts fo r

m o w i n g a n d p l o w i n g a t e a c h s i te a n d a l s o m a r k e d c o n -

T a b l e 3. R e s u l t s o f t w o - w a y A N O V A s a r e s h o w n b y y e a r f or e ac h r e s p o n se v a r i a b l e f o r d r a w d o w n p l o t s i n t h e s m a l l - s c a l e

s t u dy . F a c t o r s w e r e p l o t t y p e a n d s i t e . P l o t t y p e s c o m p a r e d i n c l u d e o n l y t r a n s p l a n te d s o i l b l o c k s a n d u n m a n i p u l a t e d c o n t r o ls .

R ~ i s t h e v a l u e f o r t h e m o d e l . ( * = s i g n i f i c a n t a t a l p h a = 0 . 0 5 ; * * = s i g n i f i c a n t a t a l p h a = 0 . 0 1 )

R e s p o n s e V a r i a b l e Y e a r R ~ S i t e p P l o t t y p e p

W e t l a n d I n d e x 1 9 9 2 0 . 9 5 1 p = 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 3 0 . 5 1 4 p = 0 . 0 1 p = 0 . 6 9

W e t l a n d S p e c i e s N u m b e r 1 9 9 2 0 . 8 1 6 p = 0 . 0 0 3 * * p < 0 . 0 0 1 *

1 9 9 3 0 . 7 7 9 p < 0 . 0 0 1 * p < 0 . 0 0 1 *

W e t l a n d S p e c i e s C o v e r 1 9 9 2 0 . 8 4 3 p = 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 3 0 . 7 0 6 p = 0 . 0 1 4 p < 0 . 0 0 1 *



428 WE TLA ND S, Volu me 17, No. 3, 1997

5

X

0

_e 4

0

~ 3

I1

<

o 2

.C

0

5

0

_c 4

01

~ 3

<

• 2

t,,,

'~ 1

0

a

a b a a

i

1 9 9 2

'

I t l l i

a ~ a ~ a

i

1 9 9 3

t t i i i

B S C L D

Tra ns p la n t P lo t Ty pe

Figure 1. Box plots of weighted average wetland index val-

ues for each transplant plot type for 1992 (after one growing

season) and 1993 (after two growing seasons) for drawdown

plots in the small-scale study. Plot types are as follows: B

= transplanted soil blocks; S = transplanted sieved soil; C

= unmanipulated controls; L = litter removed; and D =

upland soil disturbed. Plot types that share a common letter

are not significantly different based on Mann-Whitney U

tests, with alpha = 0.05. Asterisks denote outliers.

trol areas that were not disturbed. Soil was transplant-

ed with a bulldozer from the remnant wetland in the

drainage ditch in a band that extended upslope to the

edge of the proposed wetland. Mowed plots had up-

land vegetation cut but not removed, and plowed plots

had upland soil turned over, both with standard farm

equipment. Mowing treatments were established at 4

sites and plowing treatments at only 2 sites because of

problems with equipment and early autumn flooding

caused by above-average precipitation.

Wetland hydrology was restored at all five sites in

the fall of 1993. During the spring of 1994, we estab-

lished quadrats at known elevations at all sites by sur-

veying with an autolevel. Within each treatment area,

we established two replicate transects of 3 plots each

at the elevation of maximum water level and at 15 cm

and 30 cm below water level. At one very large site,

we established each treatment twice to test for consis-

tency o f results within a site and installed a set of

transects in each of the two treatment blocks. At two

of the other sites, where the soil transplant area was

at least 15-m wide, we installed an additional transect

of three plots. The total numbers of plots in each treat-

ment are shown in Table lB.

We used the wetland index described above to com-

pare the indicator status of vegetation in each plot. We

also determined the number of wetland plant species

and the total percent cover of wetland plants in each

plot. In addition, we determined the number of species

and the total cover of the plant species known to be

used as food sources by waterfowl and wetland birds,

as reported from 65 years of field and laboratory re-

search by the U.S. Fish and Wildlife Service on the

food preferences of birds (Table 2).

We compared the wetland index values, number of

wetland plant species, and total cover among treatment

types and sites with two-way ANOVAs using factors

of site and treatment type and repeated each test for

each of the three elevations in each year. We tested the

effects of elevation on wetland index values using a

two-way ANOVA with factors of treatment type and

plot elevation. We compared total cover of Typha spp.

among treatment types separately at each elevation us-

ing a one-way ANOVA. We used normal probability

plots to test assumptions of normality for data and re-

siduals. When treatment was a significant effect, we

compared pairwise combinations of treatment types

with lMkey's HSD proced ure. All statistical proc edure s

were carried out using SYSTAT (Wilkinson 1992).

RESULTS

Small-Scale Study

A natural drawdown of water level occurred during

each summer at five of the six sites. The experimental

plots at these sites had no standing water for approx-

imately two months during each growing season be-

fore reflooding in the fall. All of the data presented

are from the plots where a drawdown occurred. The

soil was never free of standing water at one site or at

the deeper plot set (-30 cm) at another site. Because

there was no drawdown at these permanently flooded

plots, there was no germination of any wetland plant

species during the course of the study, and no data

analyses are presented for these plots. Germination of

seeds in the permanently flooded plots probably was

not triggered without the fluctuating temperature re-

gime and increased oxygen levels that occur when soil

is exposed.

Transplanted Soil Blocks and Unmanipulated Con-

trols. The transplanted wetland soil blocks had sig-

nificantly lower wetland index values than the unma-

nipulated control plots in the first year but not in the

second (Table 3, Figure 1). Transplanting soil from the

renmant wetland significantly increased the number of



B r o w n & B e df o r d , W E T L A N D R E S T O R A T I O N W I T H T R A N S P L A N T E D S O I L 4 2 9

6

A M e a n N u m b e r o f W e t l a n d P l a n t S p e c i e s by T re a t m e n t

7

Block Sieved Contro l L i t te r Dis turb ed

B M e a n T o t a l C o v e r o f W e t l a n d P l a n t s b y T r e a t m e n t

50 ,

8 T

~ . so . ~ , , ,

25

Block Sieved Contro l L i t te r Dis turb ed

F i g u r e 2 . M e a n n u m b e r o f w e t l a n d p l a n t s p e c i es ( A ) a n d m e a n t o t a l p e r c e n t c o v e r o f w e t l a n d p l a n t s ( B ) i n d r a w d o w n p l o t s

o f e a c h t ype i n the s m a l l - s c a l e s t udy in e a c h ye a r , w i t h s t a nda r d de v i a t i ons . P l o t t ype s a r e a s f o l low s : B l oc k = t r a ns p l a n t e d

s o i l b l oc k ; S i e ve d = t r a ns p l a n t e d s i e ve d s o il ; Con t r o l = un ma n i pu l a t e d c on t r o l ; L i t t e r = l i t t e r r e m ove d ; a nd Di s t u r be d =

up l a nd s o i l d i s t u r be d .

w e t l a n d s p e c ie s a n d t h e a m o u n t o f c o v e r o f w e t la n d

s p e c i e s i n e a c h y e a r ( T a b l e 3 , F i g u r e 2 ) . S i t e w a s a

s i g n i fi c a n t f a c t o r i n e a c h y e a r , a n d t h e r e w e r e t w o s i g -

n i f i c a n t i n t e r a c t i o n s b e t w e e n s i t e a n d t r e a t m e n t t y p e ,

i n 1 9 9 2 f o r w e t l a n d i n d e x v a l u e ( p = 0 . 0 1 6 ), a n d i n

1 9 9 3 f o r w e t l a n d s p e c i e s c o v e r ( p < 0 . 0 0 1) .

Sieved Transplanted Soil Plots and Unmanipulated

Controls.

S i e v e d t r a n s p l a n t e d p l o t s c o n s i s t e d o f

t r a n s p l a n ts w i t h o n l y s o i l a n d n o w e t l a n d p l a n t r h i-

z o m e s . W e c o m p a r e d t h e s e p l o ts t o t h e u n m a n i p u l a t e d

c o n t r o l s a n d t o t h e t r a n s p l a n t e d s o i l b l o c k s w i t h i n

s i te s . T r a n s p l a n t i n g s i e v e d s o i l l o w e r e d t h e w e t l a n d

i n d e x v a l u e r e l a t i v e to t h e u n m a n i p u l a t e d c o n t r o l s i n

t h e f i r s t y e a r , b u t b y t h e s e c o n d y e a r , t h e v a l u e s w e r e

n o t s i g n i f i c a n t l y d i f f e r e n t ( T a b l e 4 A , F i g u r e 1 ) . I n t h e

s e c o n d y e a r, t r a n s p l a n t e d s o i l b l o c k s h a d s i g n i f ic a n t l y

m o r e w e t l a n d s p e c i e s t h a n t r a n s p l a n t e d s i e v e d s o i l ( T a -

b l e 4 A , F i g u r e 2 A ) . I n b o t h y e a r s , t h e r e w a s s i g n i f i -

c a n t l y m o r e t o t a l c o v e r f r o m w e t l a n d p l a n t s p e c i e s in

p l o t s w i t h s i e v e d s o i l t h a n i n u n m a n i p u l a t e d c o n t r o l s

( T a b le 4 A , F i g u r e 2 B ) . B o t h t h e a m o u n t o f c o v e r a n d

t h e n u m b e r o f s p e c i e s in c r e a s e d a t a l l t r e a t m e n t s i n t h e

s e c o n d y e a r r e l a t i v e t o t h e f i r s t y e a r .

Litter Removal and Disturbed Soil Plots Compared to

Unmanipulated Controls.

B o t h l it te r r e m o v a l a n d

d i s t u r b a n c e o f u p l a n d s o i l a f f e c t e d p l a n t r e s p o n s e s i n

t h e f i rs t y e a r b u t n o t i n t h e s e c o n d . I n b o t h p l o t t y p e s ,

w e t l a n d i n d e x v a l u e s w e r e s i g n i f i c a n t ly l o w e r t h a n in

u n m a n i p u l a t e d c o n t r o l s ( i n d i c a t i n g a g r e a t e r d o m i -

n a n c e o f w e t l a n d p l a n t s ) b u t o n l y i n t h e fi r st y e a r ( T a -

b l e 4 B , F i g u r e 1 ). S o il d i s t u r b a n c e a l s o i n c r e a s e d t h e

n u m b e r o f w e t la n d p l a n t s p e c i e s a n d t h e to t a l c o v e r o f

w e t l a n d p l a n t s r e l a t i v e t o u n m a n i p u l a t e d c o n t r o l s , b u t

a g a i n o n l y i n t h e f i rs t y e a r ( T a b l e 4 B , F i g u r e s 2 A a n d

B ) .


I n t h e fi rs t y e a r , s o i l t r a n s p l a n t p l o t s a t - 1 5 c m h a d

s i g n i f ic a n t l y l o w e r w e t l a n d i n d e x v a l u e s t h a n u n m a -



4 3 0 W E T L A N D S , V o l u m e 1 7, N o . 3 , 1 9 9 7

T a b l e 4. R e s u l t s o f K r u s k a l -W a l l i s o n e - w a y a n a l y s e s o f v a r i a n c e c o m p a r i n g t r a n sp l a n t t y p es a n d c o n t r o l ty p e s a t d r a w d o w n

p l o t s i n t h e s m a l l - s c a l e s t u d y fo r t h e t h r e e r e s p o n s e v a r i a b l e s : W I = W e t l a n d I n d e x ; W S N = W e t l a n d S p e c i e s N u m b e r ; W S C

= W e t l a n d S p e c i e s C o v e r . A ) C o m p a r i s o n s b e t w e e n s o i l t r a n s p la n t t y p es . B ) C o m p a r i s o n s b e t w e e n c o n t r o l ty p e s . A b b r e v i a t i o n s

f o l l o w T a b l e 1 . ( * = s i g n i f i c a n t a t a l p h a = 0 . 0 5 ; * * = s i g n i f i c a n t a t a l p h a = 0 . 0 1 )

Y e a r C o m p a r is o n W I W S N W S C

A ) S i e v e d T r a n s p l a n t e d S o i l P l o t s

1 9 92 T r a ns . B l o c k - - S i e v e d S o i l

S i e v e d S o i l - - U n m a n i p u l a te d

1 9 93 T r a ns . B l o c k s - - S i e v e d S o i l

S i e v e d S o i l - - - - - U n m a n i p u l a t e d

B ) M a n i p u l a t e d C o n t r o l P l o t s

1 9 92 L i tt e r R e m . - - U n m a n i p u l a t e d

D i s t u r b e d ~ U n m a n i p u l a t e d

1 9 93 L i tt e r R e m . - - U n m a n i p u l a t e d

D i s t u r b e d - - U n m a n i p u l a t e d

p = 0 . 4 6 5 p = 0 . 0 7 0 p = 0 . 4 2 3

p = 0 . 0 0 4 * * p = 0 . 1 2 7 p = 0 . 0 0 4 * *

p = 0 . 7 4 3 p = 0 . 0 1 3 p = 0 . 8 4 6

p = 0 . 9 4 9 p = 0 . 0 9 9 p = 0 . 0 4 8 *

p = 0 . 0 3 2 * p = 0 . 3 4 1 p = 0 . 0 2 8 *

p = 0 . 0 1 8 p = 0 . 0 4 9 * p = 0 . 0 2 8 *

p = 0 . 7 7 4 p = 0 . 3 5 9 p = 0 . 3 7 4

p = 0 . 8 3 0 p = 0 . 4 1 2 p = 0 . 2 0 0

n i p u l a t e d c o n t r o l s ( T a b l e 5 , a n d F i g u r e 3 ) . W e t l a n d

i n d e x v a l u e s a t a ll t h r e e e l e v a t i o n s s h o w e d s i m i l a r p a t -

t e r n s , w i t h n o s i g n i f i c an t in t e r a c t i o n b e t w e e n t r e a t m e n t

t y p e a n d e l e v a t i o n ( F = 1 . 6 3 4 , d f = 6 , p = 0 . 1 4 3 ) , s o

o n l y o n e e l e v a t i o n i s p r e s e n t e d g r a p h i c a l l y . S o i l t r a n s-

p l a n t s a l s o s i g n i f i c a n t l y i n c r e a s e d b o t h t h e n u m b e r o f

s p e c i e s a n d t h e c o v e r o f w e t l a n d p l a n t s a t a l l e l e v a -

t i o n s ( T a b l e 5 a n d F i g u r e 4 ) .

I n t h e s e c o n d y e ar , n o s i g n i f i c a n t d i f f e r e n c e s i n w e t -

l a n d i n d e x v a l u e s e x i s t e d a m o n g t h e t r e a t m e n t s i n

p l o t s a t - 1 5 c m ( F i g u r e 3 ) . A g a i n , t h e p a t te r n w a s s i m -

i l a r a t a ll t h r e e e l e v a t i o n s , w i t h n o s i g n i f i c a n t i n t e r -

a c t i o n b e t w e e n t r e a t m e n t t y p e a n d e l e v a t i o n ( F =

0 . 6 1 3 , d f = 6 , p = 0 . 7 1 9 ) . T h e n u m b e r o f w e t l a n d

p l a n t s p e c i e s w a s s i g n i f i c a n t l y i n c r e a s e d b y s o i l t r a n s -

p l a n t a t i o n a n d p l o w i n g ( F i g u r e 4 A ) . S o i l - t r a n s p l a n t

p l o t s h a d s i g n i f i c a n t l y m o r e s p e c i e s t h a n c o n t r o l p l o t s

a t a l l e l e v a t i o n s . P l o w e d p l o t s h a d m o r e s p e c i e s t h a n

c o n t r o l s a t t h e t w o h i g h e r e l e v a t i o n s . T h e s o i l - t r a n s -

p l a n t p l o ts a l s o h a d s i g n i f i c a n t l y g r e a t e r c o v e r t h a n

c o n t r o l s a t a l l e l e v a t i o n s ( F i g u r e 5 A ) . P l o w e d p l o t s

h a d m o r e c o v e r t h a n c o n t r o l s a t a l l e l e v a t i o n s a n d

m o r e c o v e r t h a n m o w e d p l o t s at t h e w e t l a n d e d g e . B y

t h e s e c o n d g r o w i n g s e a s o n , t h e n u m b e r s o f s p e c i e s a n d

t o t a l c o v e r a t t h e c o n t r o l p l o ts h a d i n c r e a s e d b u t w e r e

s t i l l s i g n i f i c a n t l y l o w e r t h a n t h e s o i l t r a n s p l a n t p l o t s

T a b l e 5 . R e s u l t s o f t w o - w a y A N O V A s f o r t h e l a r g e - s c a l e e x p e r i m e n t a r e s h o w n f o r e a c h r e s p o n s e v a r i a b l e a t e a c h e l e v a t io n

i n e a c h y e ar . F a c t o r s w e r e t r e a t m e n t t y p e a n d s i te , a n d R : i s th e v a l u e f o r t h e m o d e l . T r e a t m e n t s i n c l u d e d m o w i n g , p l o w i n g .

a n d t r a n s p l a n t i n g w e t l a n d s o i l , a n d c o n t r o l s w e r e n a t u r a l , u n m a n i p u l a t e d a r e a s . (* = s i g n i f i c a n t a t a l p h a = 0 . 0 5 ; * * = s i g n i f i c a n t

a t a l p h a = 0 . 0 1 )

R e s p o n s e V a r i a b l e E l e v . Y e a r R 2 S i t e p T r e a t m e n t p

W e t l a n d I n d e x E d g e 1 9 9 4 0 . 7 9 3 p = - 0 . 0 1 4 p < 0 . 0 0 1 *

1 9 9 5 0 . 7 5 1 p = 0 . 0 0 1 * p < 0 . 0 0 1 *

- 1 5 e m 1 9 9 4 0 . 5 9 5 p = 0 . 1 6 4 p < 0 . 0 0 1 *

1 9 9 5 0 . 7 9 3 p = 0 . 0 0 2 * * p = 0 . 0 4 9 *

- 3 0 c m 1 9 9 4 0 . 6 0 5 p < 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 5 0 . 7 4 6 p < 0 . 0 0 1 * p = 0 . 0 7 3

W e t l a n d S p e c i e s N u m b e r E d g e 1 9 9 4 0 . 8 2 8 p < 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 5 0 . 7 3 0 p = 0 . 0 3 5 * p < 0 . 0 0 1 *

- 1 5 c m 1 9 9 4 0 . 7 3 5 p = 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 5 0 . 8 4 7 p = 0 . 0 1 2 p < 0 . 0 0 1 *

- 3 0 c m 1 9 9 4 0 . 8 1 5 p = 0 . 0 0 2 * * p < 0 . 0 0 1 *

1 9 9 5 0 . 4 4 4 p = 0 . 1 6 0 p = 0 . 0 3 4 *

W e t l a n d S p e c i e s C o v e r E d g e 1 9 9 4 0 . 9 1 3 p = 0 . 0 0 6 * * p < 0 . 0 0 1 *

1 9 9 5 0 . 8 6 9 p < 0 . 0 0 1 * p < 0 . 0 0 1 *

1 5 c m 1 9 9 4 0 . 7 9 9 p = 0 . 1 8 1 p < 0 . 0 0 1 *

1 9 9 5 0 . 7 5 9 p < 0 . 0 0 1 * p = 0 . 0 0 2 * *

- 3 0 c m 1 9 9 4 0 . 7 01 p = 0 . 0 0 1 * p < 0 . 0 0 1 *

1 9 9 5 0 . 6 1 9 p = 0 . 0 0 1 * p - 0 . 2 0 6



Brown & Bedford, WETL AND RESTOR ATION WITH TRAN SPLA NTED SOIL 431

(Figures 4B and 5B). The number of wetland plant

species decreased in the plowed plots, which were

dominated by cattail.

The responses of the plant species used by water-

fowl and marshbirds as food sources were very similar

to the responses of wetland vegetation in general. The

soil-transplant plots had significantly greater species

number and total cover than control plots at all ele-

vations in the first year (Figures 6A and 7A). The

mow ed and plowed plots had consistently lower values

than the soil-transplant plots, but the differences were

only significant at the lowest elevation. In the second

growing season, there were significantly more wildlife

food species at the soil-transplant plots at the upper

two elevations (Figure 6B), and the average number

of species at the plowed plots had decreased. There

was significantly more cove r of wildlife food plants at

the transplant plots than at the controls (Figure 7B),

and the plowed plots had less cover than any other

treatment at the two lower elevations.

The total cover of Typha spp. was significantly in-

creased by the plowing treatment (Figure 8). At all

three elevations, the mean total cover of cattail was

significantly greater in the plowed plots than in each

of the other plot types (p<0.001 for all comparisons),

and there were no other significant differences.

DISCUSSION

Small-Scale Study

Because each restoration site had a remnant wetland

plant community in the drainage ditch and predomi-

nantly old-field species in the area that became shallow

marsh following restoration, we hypothesized that

re-establishment of hydrophytic vegetation would pro-

ceed more quickly following transplantation of wet-

land propagules from the remnant wetland soil. Our

results strongly support this hypothesis and show that

transplantation of remnant wetland soils can increase

the number and cover of wetland species in the re-

flooded areas of restoration sites. The plots with trans-

planted soil blocks had lower wetland index values,

more wetland species present, and more cove r of wet-

land plants than the unmanipulated controls in 1992.

The differences in wetland species number and cover

persisted in 1993. These results agree with similar

work reported by McKnight (1992), who found that

transplanted wetland soil was effective at establishing

wetland vegetation on disturbed sites in Texas.

The increases in wetland plant germination occurred

only in plots with a natural drawdown period. Germi-

nation of wetland plants, and in particular wetland an-

nuals, occurs most aggressively when standing water is

removed from the soil during a draw down (Gallinato and

a a a b b

1 9 9 4

5

o 3

>

< [ 2

a a a a

1 9 9 5

C M P S T

T r e a tm e n t y p e

Figure 3. Box plots of weighted average wetland index val-

ues for plots at -15cm in each large-scale experiment plot

type in 1994 and 1995. Plot types are as follows: C = un-

manipulated control; M = mowed; P = plowed; ST = wet-

land soil transplanted. Plot types that do not share a common

letter are significantly different at alpha = 0.05. Asterisks

denote outliers and circles extreme outliers.

van der Valk 1986). Drawdowns have been shown to

have a significant influence on vegetation communities

in freshwater wetlands (Harris and Marshall 1963, van

der Valk and Davis 1978). The methods studied here

would be most effective at increasing wetland plant es-

tablishment in areas of restored wetland basins that have

a natural drawdown at least in some years. We expect

that over a longer time period, wetland plants will even-

tually become established at lower elevations as well, but

the study period was too short to observe this aspect of

plant co mmuni ty restoration.

Many of the other effects that were significant in

the first year did not persist into the second year. In

the first year, manipulated control plots were similar

to transplanted soil blocks, with similar wetland index

values and amounts of cover of wetland plants. After

the first year of the study, it seemed that these methods

might be considered as alternatives to actual trans-

plantation of wetland soil. By the second year, how-

ever, manipulated upland plots without soil amend-

ments showed no improvement in any of the response

variables relative to the unmanipulated controls. After

two years of flooding, enough wetland plant establish-



4 3 2 W E T L A N D S , V o l u m e 1 7, N o . 3 , 1 99 7

1 4

A M e a n N u m b e r o f W e t l a n d P l a n t S p e c i e s in 1 9 9 4

a a a b a a b b c c a a b c

1 2

a T

0

-3 0 -1 5 E d g e

[ ] Contro l

13

M o w e d

E] P l o w e d

[ ] S o i l T ra n s p l a n t

1 4

12

1 0

6

E

B M e a n N u m b e r o f W e t l a n d P l a n t S p e c i e s i n 1 9 9 5

a a a b a a b b c a a a b

i l i _

3 0 -1 5 E d g e

I I C o n t ro l

- - [ ] M o w e d

[ ]

P l o w e d

I ~ S o i l T ra n s p l a n t

Fi gu r e 4 . M e a n nu m be r o f we t l a nd p l a n t s pe c i e s in 1994 ( A ) a nd 1995 ( B) in p l o t t ype s i n t he l a r ge - s c a l e e xpe r i me n t , w i t h

s t a nda r d de v i a t i ons . T r e a t me n t s w i t h i n e a c h e l e va t i on t ha t do no t s ha r e a c ommon l e t t e r a r e s i gn i f i c a n t l y d i f f e r e n t a t a l pha =

0.05.

m e n t h a d o c c u r r e d i n t h e u n m a n i p u l a t e d c o n t r o l s t o

m a k e t h e m s i m i l a r t o t h e l i t t e r - r e m o v a l a n d s o i l - d i s -

t u r b a n c e p l o t s . T h u s , i t s e e m s t h a t w h i l e t e m p o r a r y

e f f e c t s c a n b e p r o d u c e d b y r e m o v i n g l it te r o r d i s t u r b -

i n g t h e u p l a n d s o il , o n l y t h e a d d i t i o n o f s o i l f r o m t h e

r e m n a n t w e t l a n d h a s l o n g e r t e r m e f f e c t s o n t h e w e t -

l a n d p l a n t c o m m u n i t y t h a t d e v e l o p s . T r a n s p l a n t a t i o n

o f r e m n a n t w e t l a n d s o il s e e m s t o b e t h e m o s t e f f e c t iv e

t e c h n i q u e f o r i n c r e a s i n g t h e re - e s t a b l i s h m e n t o f w e t -

l a n d p l a n t s r e l a t i v e t o n a t u r a l r e c o l o n i z a t i o n .

T h e r e w e r e n o d i f f e r e n c e s b e t w e e n p l o t s w i t h t r a n s -

p l a n t e d s o i l a n d p l o t s w i t h t r a n s p l a n t e d s o i l a n d r h i z o m e s

i n t h e f ir s t y e a r , w i t h b o t h t y p e s s h o w i n g s i g n i f i c a n t i n -

c r e a s e s i n n u m b e r a n d t o t a l c o v e r o f w e t l a n d p l a n t s p e -

c i e s c o m p a r e d t o u n m a n i p u l a t e d c o n t r o ls . T h e s e re s u l t s

s u g g e s t t h a t , d u r i n g t h e f i rs t g r o w i n g s e a s o n f o l l o w i n g

t r a n s p l a n t a t i o n , t h e r e i s n o p r a c t i c a l d i f f e r e n c e i n t h e

r e - e s t a b l i s h m e n t o f w e t l a n d p l a n t s t h a t r e s u l ts f r o m t h e s e

t w o m e t h o d s . H o w e v e r , i n t h e s e c o n d y e a r , t h e p lo t s w i t h

t r a n s p l a n t e d r h i z o m e s h a d s i g n i f i c a n tl y g r e a t e r n u m b e r s

o f w e t l a n d s p e c i e s t h a n t h o s e w i t h o n l y s i e v e d s o i l. T h i s

s u g g e s t s t h a t o v e r a l o n g e r te r m , r h i z o m a t o u s s p e c i e s c a n

b e c o m e e s t a b l i s h e d a n d i n c r e a s e t h e s p e c i e s d e n s i t y i n

p l o t s w h e r e t h e y a r e tr a n s p l a n t ed . L o n g - t e r m s t o r a g e o f

s o i l b e f o r e t r a n s p l a n t a t i o n a t r e s t o r a t i o n s i t e s c o u l d r e -

d u c e t h e n u m b e r o f v i a b le r h i z o m e s a n d t h e r e f o r e r e d u c e

t h e n u m b e r o f s u c c e s sf u l ly e s t a b li s h e d s p e c i e s. T h i s m a y

b e s i g n i f i c a n t s i n c e r h i z o m e s a r e a s s o c i a t e d w i t h p e r e n -

n i a l e m e r g e n t s t h a t a r e i m p o r t a n t f o r r e s t o r a t io n o f a v i -

f a u n a f o o d a n d c o v e r ( M a r t i n e t al . 1 9 5 1 , P a y n e 1 9 9 2 ).

T h e r e s u l ts f r o m t h e s m a l l - s c a le e x p e r i m e n t s h o w

t h at t h e t h re e r e s p o n s e v a r i a b l e s o f n u m b e r o f w e t l a n d

p l a n t s p e c i e s , a m o u n t o f c o v e r , a n d w e t l a n d i n d e x a ll

r e f l ec t d i f f e r e n t c h a r a c t e r i s ti c s o f t h e r e s t o r e d p l a n t

c o m m u n i t y . I n th e s e c o n d y e a r o f t h e s t u d y , w e t l a n d

p l a n t s a t t r a n s p l a n t e d s o i l b l o c k s h a d g r e a t e r c o v e r

t h a n a t u n m a n i p u l a t e d c o n t r o l s , b u t th e w e t l a n d i n d e x

v a l u e s f o r t h e t w o t y p e s o f p lo t s d i d n o t d i f fe r . T h i s

w a s t r u e e v e n t h o u g h t h e w e t l a n d i n d e x v a l u e s w e r e

c a l c u l a t e d u s i n g t h e r e l a t iv e c o v e r o f e a c h s p e c i e s .

W i t h i n t w o g r o w i n g s e a s o n s a f t e r w e t l a n d h y d r o l o g y

w a s r e s t o r e d , o n l y p l a n t s p e c i e s t h a t c o u l d t o l e r a t e s a t-



B r o w n & B e df o r d , W E T L A N D R E S T O R A T I O N W I T H T R A N S P L A N T E D S O I L 4 33

E

n

I--

3 0 0

2 5 0

2 0 0

1 5 0

1 0 0

5 0

0

A M e a n T o t a l C o v e r o f W e t l a n d P l a n t s i n 1 9 9 4

a a b bc c a b b c a a b c

I

- 3 0 - 1 5 E d g e

• C o n t ro l

IR M o w e d

[ ] P l o w e d

[ ] S o i l T r a n s p l a n t

3 0 0

2 5 0

> 2 0 0

B M e a n T o t a l C o v e r o f W e t l a n d P l a n t s i n 1 9 9 5

a a a a a a b a b b a a a b b

O t i i

1 5 0

~ 1 0 0

5 0

0

-3 0 -1 5 E d g e

• C o n t ro l

• M o w e d

r l P I o w e d

[ ] S o i l T ra n s p l a n t

Fi gu r e 5 . M e a n t o t a l pe r c e n t c ov e r o f we t l a nd p l a n t s i n t he la r ge - s c a l e e xpe r i m e n t i n 1994 ( A ) a nd 1995 ( B) , w i t h p l o t

l a you t f o l l owi ng F i gu r e 4 .

u r a t e d s o il s p e r si s t e d , s o t h e i n d e x v a l u e s o f t h e v e g -

e t a t i o n p r e s e n t w e r e u n i f o r m l y l o w a c r o s s a l l t r e a t m e n t

t y p e s . A t t h e s a m e t i m e , t h e t r a n s p l a n t e d s o i l b l o c k s

h a d s i g n i f i c a n t ly m o r e c o v e r o f w e t l a n d p l a n t s a n d

m o r e p l a n t s p e c i e s . T h e i n d e x v a l u e s r e f l e c te d th e

c h a n g e s i n v e g e t a t i o n t h a t o c c u r r e d i n t h e fi rs t g r o w i n g

s e a s o n f o l l o w i n g r e s t o r a t i o n o f h y d r o l o g y . H o w e v e r ,

t h e i n d e x v a l u e d i d n o t r e f l e c t t h e d i f f e r e n c e s i n t h e

w e t l a n d p l a n t c o m m u n i t y t h a t p e r s i s t e d a ft e r t h e m a -

j o r i t y o f s u r v i v i n g p l a n t s w e r e w e t l a n d s p e c i e s . A n a l -

y s i s o f re s t o re d p l a n t c o m m u n i t i e s i n v o l v i n g c o m p a r -

i s o n w i t h r e f e r e n c e w e t l a n d s s h o u l d i n c l u d e m e a s u r e -

m e n t o f s p e c i e s ri c h n e ss a n d c o v e r b e c a u s e i n d e x v a l -

u e s c a n n o t d i s c r i m i n a t e i m p o r t a n t d i f f e r e n c e s i n

c o m m u n i t y s t r u c t u r e a f t e r t h e in i t ia l r e a d j u s t m e n t to

w e t l a n d c o n d i t i o n s .


T h e l a r g e - s c a l e s t u d y c o n f i r m e d t h a t t h e i n c r e a s e s

i n w e t l a n d p l a n t r e s p o n s e s o b s e r v e d i n t h e s m a l l - s c a l e

s t u d y c o u l d b e a c h i e v e d a t t h e s c a l e o f w h o l e s it e s.

T h e l a r g e - s c a l e r e s u l t s s h o w e d t h a t s o i l t r a n s p l a n t a t i o n

c a n i n c r e a se t h e n u m b e r a n d t h e c o v e r o f w e t l a n d p l a n t

s p e c i e s a t r e s t o r e d w e t l a n d s a n d t h a t t h e t e c h n i q u e c a n

b e a p p l i e d a t t h e s c a l e o f w h o l e s i te s d u r i n g i n i t ia l

r e s to r a t io n w o r k . I m p r o v e m e n t i n a v i f a u n a h a b i ta t r e-

q u i re s i n c r e a s e s in t h e n u m b e r a n d c o v e r o f w e t l a n d

p l a n t s t h a t a re u s e d a s f o o d s o u r c e s . T h e t r a n s p l a n t

t r e a tm e n t s s i gn i fi c a nt ly i m p r o v e d b o t h t h e n u m b e r o f

s p e c i es a n d t h e a m o u n t o f c o v e r o f f o o d p l an t s . T h i s

t r e a t m e n t c o u l d s i g n i f i c a n t ly i m p r o v e th e h a b i t a t v a l u e

o f r e s t o r e d s i t e s f o r a v i f a u n a , p a r t i c u l a r l y i n t h e f i rs t

f e w y e a r s . S i e g l e y e t a l. ( 1 9 8 8 ) f o u n d s i m i l a r r e s u l t s

f o r m a r s h e s c r e a t e d u s i n g d r e d g e s p o il s o n L a k e E r i e

t h a t c o n t a i n e d w e t l a n d p l a n t s e e d s , w h i c h s h o w e d i n -

c r e a s e s in w i l d l if e f o o d p l a n t d e v e l o p m e n t w h e n s e e d -

r i c h s p o i l s w e r e u s e d .

H o w e v e r , w e d o n o t k n o w h o w l o n g th e e f f e c t o f

s o i l t ra n s p l a n t s w i l l p e rs i s t , a n d s h o r t - t e r m e s t a b l i s h -

m e n t p a t te r n s m a y n o t p r e d i c t l o n g - t e r m p l a n t c o m -

m u n i t i e s ( W e i h e r e t al . 1 9 9 6 ). I t s e e m s l i k e l y f r o m t h e



4 3 4 W E T L A N D S , V o l u m e 1 7, N o . 3 , 1 9 97

z

A M e a n N u m b e r o f W i l d l i f e F o o d P l a n ts in 1 9 94

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Fi gu r e 6 . M e a n nu mb e r o f w i l d l i f e food p l a n t s pe c i e s in the l a r ge - s c a l e e xpe r i me n t i n 1994 ( A ) a nd 1995 ( B) , w i t h p l o t


r e s u l t s a t o t h e r r e s t o r e d w e t l a n d s i n o u r s t u d y t h a t t h e

u n t r e a t e d a r e a s w i l l d e v e l o p s i m i l a r s p e c i e s r i c h n e s s

a n d c o v e r o f h e r b a c e o u s w e t l a n d p l a n t s o v e r a p p r o x -

i m a t e l y t h r ee y e a r s ( B r o w n 1 9 95 ) . I f t h e im p r o v e m e n t

f r o m s o i l t r a n s p l a n t s d o e s n o t p e r s i s t l o n g e r t h a n t h e

f i rs t f e w g r o w i n g s e a s o n s , t h e n t h e t e c h n i q u e w o u l d

b e u n n e c e s s a r y a s a g e n e r a l p r a c t i c e .

T h e m o s t i m p o r t a n t u s e o f th i s te c h n i q u e m a y b e t o

c o n t r o l c a t t a il i n a r e a s w h e r e s o i l h a s b e e n d i s t u r b e d

d u r i n g c o n s t r u c t i o n . S o i l d i s tu r b a n c e i n t h e p l o w e d

t r e a t m e n t s h a d t h e p r e d i c t e d e f f e c t o f i n c r e a s i n g c o l -

o n i z a t i o n b y s e e d l i n g s o f Typha s p p . , a n d w e m e a s u r e d

s i g n i f ic a n t l y l o w e r n u m b e r s a n d c o v e r o f w i l d l i f e f o o d

p l a n t s i n t h e s e a r e a s . T h i s h a s b e e n a s i g n i f i c a n t p r o b -

l e m a t o t h e r r e s t o r a t i o n s i t e s i n o u r s t u d y , p a r t i c u l a r l y

i n t h e a r e a w h e r e d i k e c o n s t r u c t i o n d i s t u r b s t h e s o i l

( B r o w n 1 9 9 5 ). M o n o t y p i c s t a n d s o f c a t ta i l r e d u c e v e g -

e t a t i o n d i v e r s i t y a n d t h e r a t i o o f o p e n w a t e r t o v e g e -

t a t i o n , a n d , t h e r e f o r e , t h e h a b i t a t v a l u e o f t h e s i te ( W e l -

l e r 1 9 7 5, K a m i n s k i a n d P r i n c e 1 9 8 1 , M u r k i n e t a l.

1 9 8 2 ). R e d u c t i o n o f ca t t a il t h r o u g h t h e u s e o f

c o a r s e - t e x t u r e d s u b s t r a te s , a s s u g g e s t e d b y W e i h e r e t

a l . ( 19 9 6 ), m a y b e e f f e c t i v e a t c r e a t e d w e t l a n d s b u t

w o u l d b e c o s t - p r o h i b i t i v e f o r l a r g e - s c a l e r e s t o r a t io n

p r o j e c t s . T r a n s p l a n t in g r e m n a n t w e t l a n d s o i l i n t o th e

a r e a d i s t u r b e d d u r i n g d i k e c o n s t r u c t i o n s h o u l d s u b -

s t an t ia l ly i m p r o v e t h e e s t a b li s h m e n t o f o t h e r m o r e

v a l u a b l e w e t l a n d p l a n t s p e c i e s i n t h e s e a r e a s a n d l i m i t

p o p u l a t i o n s o f c a t ta i l. V i v i a n - S m i t h a n d H a n d e l ( 1 9 9 6 )

a l so r e c o m m e n d t h e u s e o f i m p o r t e d s o il s to i m p r o v e

w e t l a n d r e s t o r a t io n s i t es . E v e n t h o u g h t h i s t e c h n i q u e

i n c r e a s e s t h e i n i ti a l c o s t o f c o n s t r u c t i o n , i t m a y p r e v e n t

t h e n e e d f o r c o s tl y i n te n s i v e m a n a g e m e n t t e c h n i q u e s

t o c o n t ro l c a t ta i l a n d c a n b e a c c o m p l i s h e d w h i l e c o n -

s t r u c t io n e q u i p m e n t i s o n s it e .

T h e e f f e c t o f t r a n s p l a n t e d w e t l a n d s o i l s o n p l a n t re -

c r u i t m e n t i n a r e s t o r e d w e t l a n d w i l l v a r y d e p e n d i n g

o n t h e c o m p o s i t i o n o f t he s e e d b a n k . W e s t u d i e d t h e

s e e d b a n k i n t h e r e m n a n t w e t l a n d s o i l s b e f o r e u n d e r -

t a k i n g t h i s e x p e r i m e n t a n d f o u n d t h a t t h e y c o n t a i n e d

a v a r i e t y o f w e t l a n d p l a n t s p e c i e s u s e f u l f o r h a b i t a t

i m p r o v e m e n t ( B r o w n 1 9 9 5) . I n ad d i ti o n , t he s e e d b a n k



B r o w n & B e df o r d , W E T L A N D R E S T O R A T I O N W I T H T R A N S P L A N T E D S O I L 4 3 5

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Fi gu r e 7 . M e a n t o t a l pe r c e n t c ov e r o f w i l d l i f e f ood p l a n t s i n the l a r ge - s c a l e e xpe r i me n t in 1994 ( A ) a nd 1995 ( B) , w i t h p l o t


l a c k e d u n d e s i r a b l e i n v a s i v e s p e c i e s s u c h a s p u r p l e

l o o s e s t r i f e

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L . ) . T r a n s p l a n t i n g r e m -

n a n t w e t l a n d s o i l i s p r u d e n t o n l y w h e n t h e c o m p o s i -

t i o n o f t h e s e e d b a n k i s k n o w n a n d i s c o n s i s t e n t w i t h

m a n a g e m e n t g o a l s ( P e d e r s o n a n d v a n d e r V a l k 1 9 84 ,

v a n d e r V a l k e t a l. 1 9 9 2 ) . F o r e x a m p l e , t r a n s p l a n t i n g

r e m n a n t w e t l a n d s o i l f r o m a r e a s d o m i n a t e d b y c a t ta il ,

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4 3 6 W E T L A N D S , V o l u m e 1 7 , N o . 3 , 1 9 9 7

l a r g e n u m b e r s o f c a t t a i l s e e d s , w o u l d p r o b a b l y n o t i n -

c r e a s e w e t l a n d p l a n t d i v e r s i t y a s e f f e c t i v e l y .

T h e w a t e r l e v e l i s a s i g n i f i c a n t f a c t o r c o n t r o l l i n g

s e e d g e r m i n a t i o n a n d , t h e r e f o r e , p l a n t e s t a b l i s h m e n t i n

a w e t l a n d ( K e d d y a n d E l l i s 1 9 8 5 , M o o r e a n d K e d d y

1 9 8 8 ) . I n o u r s t u d y , e s t a b l i s h m e n t o f w e t l a n d p l a n t s

d e c r e a s e d s t e a d i l y w i t h i n c r e a s i n g d e p t h r e l a t i v e t o

m a x i m u m h i g h w a t e r . W e o b s e r v e d l i t tl e e s t a b l i s h m e n t

a t w a t e r d e p t h s g r e a t e r t h a n 4 5 c m e v e n t h o u g h t h e

t r e a t m e n t s e x t e n d e d a c r o s s th e e n t i r e w e t l a n d b a s i n ,

T h i s s u g g e s t s t h a t t r a n s p l a n t a t i o n o f w e t l a n d s o i l

s h o u l d b e c o n c e n t r a t e d a t s h a l l o w e r e l e v a t i o n s n e a r t h e

p r o p o s e d h i g h w a t e r l e v e l o f r e s t o r e d s i t e s .

R e l o c a t i o n o f w e t l a n d s o i l s h o u l d b e c o n s i d e r e d a s

a p o t e n t i a l l y u s e f u l r e s t o r a t i o n t e c h n i q u e w h e n e v e r

t h e r e i s r e m n a n t w e t l a n d s o i l p r e s e n t o n t h e s i te . S e e d

b a n k a s s a y s s h o u l d b e c o n d u c t e d t o e n s u r e t h a t t h e

s p e c i e s c o m p o s i t i o n i s a p p r o p r i a t e f o r t h e g o a l s o f t h e

r e s t o r a t i o n p r o j e c t . T h e u s e o f r e m n a n t s e e d b a n k s p r o -

v i d e s a r e l a t i v e l y l o w - c o s t s o u r c e o f w e t l a n d p l a n t

p r o p a g u l e s t h a t c a n s i g n i f i c a n t l y i n c r e a s e t h e r e s t o r a -

t i o n o f p l a n t c o m m u n i t i e s a t d r a i n e d w e t l a n d s . T h e

t e c h n i q u e i s p a r t i c u l a r l y u s e f u l t o p r e v e n t t h e e s t a b -

l i s h m e n t o f m o n o t y p i c c a t t a i l s t a n d s i n a r e a s w h e r e

s o i l h a s b e e n d i s t u r b e d b y d i k e c o n s t r u c t i o n .

A C K N O W L E D G M E N T S

T h i s r e s e a r c h w a s s u p p o r t e d b y R e g i o n 5 o f t h e U . S .

F i s h a n d W i l d l i f e S e r v i c e w i t h a d d i t i o n a l f i n a n c i a l

s u p p o r t f r o m D u c k s U n l i m i t e d , I n c . , t h e N a t i o n a l F i s h

a n d W i l d l i f e F o u n d a t i o n , t h e E d n a B a i l e y S u s s m a n

F u n d , a n d C o r n e l l U n i v e r s i t y . O u r f i e l d a s s i s t a n t s , in -

c l u d i n g B i l l Y a m e l l , M e t t a M c G a r v e y , a n d C h r i s t i a n

O t t k e , w o r k e d t i r e l e s s l y , c h e e r f u l l y , a n d o f t e n i n d e -

p e n d e n t l y a n d c o n t r i b u t e d i m m e a s u r a b l y t o t h e p r o j -

e c t . K a r e n P o i a n i p r o v i d e d a d v i c e a n d m o r a l s u p p o r t

t h r o u g h o u t t h e p r o j e c t a n d p r o v i d e d d e t a i l e d c o m -

m e n t s o n t h e m a n u s c r i p t . T h e m a n u s c r i p t a l s o b e n e -

f i te d f r o m t h e c o m m e n t s a n d s u g g e s t i o n s o f t w o a n o n -

y m o u s r e v i e w e r s .

L I T E R A T U R E C I T E D

B a l l , J . P . 1 9 9 0 . I n f l u e n c e o f su b se q u e n t f l o o d in g d e p th o n c a t t a i l

c o n t r o l b y b u r n i n g a n d m o w i n g . J o u r n a l o f A q u a t i c P l a n t M a n -

a g e m e n t 2 8 : 3 2 - 3 6 .

B e u l e , J . D . 1 9 7 9 . C o n t r o l a n d m a n a g e m e n t o f c a t ta i l s i n s o u th -

e a s t e r n W i s c o n s i n w e t l a n d s . D e p a r t m e n t o f N a t u r a l R e s o u r c e s ,

M a d i s o n , W I , U S A . T e c h n i c a l B u l le t i n N o . 1 1 2

B r o w n , S . C . 1 9 9 5 . W e t l a n d r e s to r a t i o n : Fa c to r s c o n t r o l l i n g p l a n t

c o m m u n i t y r e s p o n s e a n d a v i f a u n a l h a b i t a t v a l u e . P h . D . D i s s e r t a -

t i o n . C o r n e l l Un iv e r s i t y , I t h a c a , NY, USA .

C o m e s , R . D . a n d A . D . K e l l y. 1 9 8 9. C o n t r o l o f c o m m o n c a t t a i l

w i t h p o s t e m e r g e n c e h e r b i c i d e s . J o u r n a l o f A q u a t i c P l a n t M a n a g e -

m e n t 2 7 : 2 0 - 2 3 .

C o wa r d in , L . M . , V . C a r t e r . E C . Go le t , a n d E . T . L a r o e . 1 9 7 9 .

C l a s s i f i c a t i o n o f w e t l a n d s a n d d e e p w a t e r h a b i t a t s o f t h e U n i t e d

S t a t e s . U . S . F i s h a n d W i l d l i f e S e r v i c e , W a s h i n g t o n , D C , U S A .

F W S / O B S - 7 9 / 3 1 .

D u n n , W . J . a n d G . R . B e s t . 1 9 8 4 . E n h a n c i n g e c o l o g i c a l s u c c e s s i o n .

5 . S e e d b a n k s u r v e y o f s o m e F l o r i d a m a r s h e s a n d r o l e o f s e e d

b a n k s i n m a r s h r e c l a m a t i o n , p . 3 6 5 - 3 7 0 .

In

D . H . G r a v e s ( e d . )

P r o c e e d i n g s , 1 9 8 3 S y m p o s i u m o n S u r f a c e M i n i n g , H y d r o l o g y ,

S e d i m e n t o l o g y , a n d R e c l a m a t i o n . O f f i c e o f E n g i n e e r i n g S e r v i c e s ,

U n i v e r s i t y o f K e n t u c k y , L e x i n g t o n , K Y , U S A .

E i c h e r , A . L . 1 98 8. S o i l - v e g e t a t i o n c o r r e l a t i o n s i n w e t l a n d s a n d a d -

j a c e n t u p l a n d s o f t h e S a n - F r a n c i s c o B a y e s t u a r y . C a l i f o rn i a . U . S .

F i s h a n d W i l d l i f e S e r v i c e , W a s h i n g t o n , D C , U S A . B i o l o g i c a l R e -

p o r t 8 8 ( 2 1 ) .

G a l a t o w i t s c h , S . M . a n d A . G . v a n d e r V a l k . 1 9 9 6 . T h e v e g e t a t i o n

o f r e s t o r e d a n d n a t u r a l p r a i r i e w e t l a n d s . E c o l o g i c a l A p p l i c a t i o n s

6 : 1 0 2 - 1 1 2 .

G a l l i n a t o , M , I . a n d A . G . v a n d e r V a l k . 1 9 8 6 . S e e d g e r m i n a t i o n

t r a it s o f a n n u a l s a n d e m e r g e n t s r e c r u i t e d d u r i n g d r a w d o w n s i n th e

d e l t a m a r s h , M a n i t o b a , C a n a d a . A q u a t i c B o t a n y 2 6 : 8 9 - 1 0 2 .

G l e a s o n , H . A . a n d A . C r o n q u i s t . 1 9 9 1 . M a n u a l o f V a s c u l a r P l a n t s

o f N o r t h e a s t e r n U n i t e d S t a t e s a n d A d j a c e n t C a n a d a . N e w Y o r k

B o t a n i c a l G a r d e n , B r o n x , N e w Y o r k , N Y , U S A .

Ha r r i s , S . W . a n d W . H . M a r sh a l l , 1 9 6 3 . Ec o lo g y o f wa t e r l e v e l

m a n i p u l a t i o n s o n a n o r t h e r n m a r s h . E c o l o g y 4 4 : 3 3 1 - 3 4 3 .

K a m i n s k i , R . M . a n d H . H . P r i n c e . 1 9 8 1 . D a b b l i n g d u c k a n d a q u a t i c

m a c r o i n v e r t e b r a t e r e s p o n s e s t o m a n i p u l a t e d w e t l a n d h a b i t a t . Jo u r -

n a l o f W i l d l i f e M a n a g e m e n t 4 5 : 1 - 1 5 .

Ke d d y , P . A . a n d T . H . E l l i s . 1 9 8 5 . S e e d l in g r e c r u i tm e n t o f 1 1 we t -

l a n d p l a n t s p e c i e s a l o n g a w a t e r l e v e l g r a d i e n t : S h a r e d o r d i s t i n c t

r e s p o n s e s ? C a n a d i a n J o u r n a l o f B o t a n y 6 3 : 1 8 7 6 - 1 8 7 9 .

L e w i s , R . R . 1 9 8 9. W e t l a n d r e s t o r a t i o n / c r e a t i o n / e n h a n c e m e n t t e r-

m i n o l o g y : S u g g e s t i o n s f o r s t a n d a r d i z a t io n , p . 1 - 8 .

In

J . A . Ku s l e r

a n d M , E . K e n t u l a ( e d s . ) W e t l a n d C r e a t i o n a n d R e s t o r a t i o n : t h e

S t a t u s o f t h e S c ie n c e . U . S , E P A , W a s h i n g t o n , D C , U S A . 6 0 0 / 3 -

8 9 /0 3 8 a .

M a l l i k , A . U . a n d R . W . W e i n . 1 9 8 6 . R e s p o n s e o f a

typha

m a r s h

c o m m u n i t y t o d r a i n i n g , f lo o d i n g , a n d s e a s o n a l b u r n i n g . C a n a d i a n

J o u r n a l o f B o t a n y 6 4 : 2 1 3 6 - 2 1 4 3 .

M a r t i n , A . C . , H . S . Z i m , a n d A . L . N e l s o n , 1 9 5 1 . A m e r i c a n W i l d l i f e

a n d P l a n t s : A G u i d e t o W i l d l i f e F o o d H a b i t s . D o v e r P u b l i c a t i o n s ,

N e w Y o r k , N Y , U S A .

M c K n i g h t , S . K . 1 9 9 2 . T r a n s p l a n t e d s e e d b a n k r e s p o n s e t o d r a w -

d o w n t i m e i n a c r e a t e d w e t l a n d i n e a s t T e x a s . W e t l a n d s 1 2 : 7 9 -

90 .

M i c h e n e r , M . C . 1 9 8 3 . W e t l a n d s i t e i n d e x f o r s u m m a r i z i n g b o t a n i c a l

s t u d ie s . W e t l a n d s 3 : 1 8 0 - 1 9 1 .

M o o r e , D . R . J . a n d P . A . Ke d d y . 1 9 8 8 . E f f e c t s o f a wa t e r - d e p th

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o f B o t a n y 6 6 : 5 4 8 - 5 5 2 .

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Wetland Restoration With Plants