WETLANDS, Vol. 17, No. 3, September 1997, pp.

424-437
1997, The Society of Wetland Scientists

RESTORATION OF WETLAND VEGETATION WITH TRANSPLANTED

WETLAND SOIL: AN EXPERIMENTAL STUDY
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, N Y 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:

Restoration of drained wetlands requires the re-establishment of a native wetland plant community. This can be difficult in areas where long-term drainage has eliminated wetland vegetation and significantly reduced the number of viable wetland plant seeds in the seed bank. This study of U.S. Fish and
Wildlife Service wetland restoration sites in northern New York examines the effectiveness of transplanting
wetland soil from small remnant wetlands in the drainage ditches to the area that becomes shallow marsh
following reflooding. The results of two experiments are reported, including a small-scale study of transplantation techniques using small plots with treatments and controls established by hand, and a large-scale
application of soil transplantation and site-preparation techniques using heavy equipment to establish large
plots across entire wetland basins. In the small-scale study, the transplant plots had significantly lower
wetland index values, indicating greater dominance of wetland plants, after one growing season but not after
two. Transplant plots also had more wetland plant species and more wetland plant cover than natural control
plots, and these differences persisted through the second growing season. Litter removal and soil disturbance
also lowered the wetland index values and increased wetland plant species number and cover, but only for
the first growing season. In the large-scale study, soil transplantation significantly increased both the number
of species and the amount of cover of wetland plants and of plants valuable as wildlife food sources. Mowing
and plowing treatments increased wetland plant establishment, but much less than soil transplantation, and
plowing significantly increased the establishment of cattail (Typha spp.), an undesirable invasive species in
small wildlife marshes. Soil transplantation should be a particularly effective technique for improving wetland
plant establishment and limiting cattail encroachment in areas disturbed by dike construction,

Key Words:

wetland restoration, wetland plants, soil amendments, soil transplantation, New York

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 in r e s t o r e d w e t lands.
C a t t a i l i n v a s i o n is 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 the ratio o f o p e n w a t e r to v e g e t a t i o n ,
r e d u c e the 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 l u e o f the site ( W e l l e r 1975, K a m i n s k i
a n d P r i n c e 1981, M u r k i n et al. 1982). In a d d i t i o n , cattails are v e r y difficult to c o n t r o l o n c e t h e y are e s t a b lished, even with intensive management techniques
l i k e cutting, b u r n i n g , a n d the use o f h e r b i c i d e s ( B e u l e
1979, M a l l i k a n d W e i n 1986, C o m e s a n d K e l l y 1989,
B a l l 1990).
G o v e r n m e n t a l a g e n c i e s i n v o l v e d in r e s t o r a t i o n (sensu L e w i s 1989) 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 that i n c r e a s e the l i k e l i h o o d o f e s t a b l i s h i n g di-

INTRODUCTION
T h r o u g h o u t the U n i t e d States, 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 that 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 activities b e r e p l a c e d w i t h c r e a t e d o r r e s t o r e d sites. H o w ever, f e w studies 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 the eff 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 to r e c r e a t e natural w e t l a n d s . W e f o u n d in an 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 the 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 1995) that cattail (Typha spp.) o f t e n d o m i n a t e s
a r e a s o f the sites w h e r e c o n s t r u c t i o n a c t i v i t i e s disturb
the soil d u r i n g r e s t o r a t i o n . O d u m (1988), R e i n a r t z a n d
W a r n e (1993), 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
the N a t i o n a l R e s e a r c h C o u n c i l (1992) also r e p o r t the
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 cattail 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 to p r e c l u d e e s t a b 424

B r o w n & Bedford, 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
verse plant communities that resemble natural wetlands (National Research Council 1992). If increasing
avifauna habitat is a m o n g the restoration goals, the
techniques must establish plant communities that enhance habitat value for birds. Furthermore, the techniques must have the characteristic that they can be
applied to m a n y wetlands at the scale o f entire restoration sites and without excessive cost to the agency
or landowner.
Numerous studies of natural wetlands have tied the
maintenance of a diverse flora to the presence o f a diverse seed bank that responds to changing water levels
(van der Valk and Davis 1978, van der Valk 1981, Pederson and van der Valk 1984, Keddy and Ellis 1985).
Seed banks will not contribute to restoration o f wetland
plant communities, however, if they have been significantly depleted by long-term disturbance o f the former
wetland site. After m a n y years of drainage, for example,
the seed bank in a former wetland can b e c o m e relatively depauperate of wetland species (Weinhold and
van der Valk 1989, Brown 1995, Galatowitsch and van
der Valk 1996). This loss of viable seeds may slow or
prevent recovery o f wetland vegetation following restoration of wetland water levels. Desirable species m a y
not re-establish before a more aggressive species preempts the site (Reinartz and Warne 1993).
On sites drained with ditches, however, a remnant
wetland plant c o m m u n i t y often exists in the poorly
drained soil adjacent to the ditch. We hypothesized
that, if these remnant soils could be transplanted to the
intended restoration area, their relatively rich seed
banks would produce greater species richness and cover o f wetland plants following restoration of wetland
hydrology than the seed banks of areas that had been
effectively drained (Dunn and Best 1984). We further
hypothesized that the technique o f m o v i n g remnant
wetland soils was one that could be i m p l e m e n t e d on
a scale large enough to affect entire restoration sites.
Our goal was to determine if transplanting remnant
wetland soil could be an effective technique for recreating a wetland plant c o m m u n i t y at higher elevations
where drainage was m o r e complete. We conducted our
experiments at two scales, a small-scale study using
p l o t s with treatments established b y hand, and a largescale study using construction equipment to establish
treatment areas across entire wetland basins. Because
the goal of the restoration p r o g r a m we studied was to
increase avifauna habitat, we were particularly interested in the responses of wetland plants k n o w n to be
used as food sources by waterfowl and marsh birds.
Our experiments were designed to address the following major questions: 1) can the n u m b e r of wetland
plant species that germinate and establish be increased
by transplanting soil f r o m remnant wetland areas; 2)
does the inclusion of wetland plant rhizomes in the

425

transplanted soil affect the species richness and abundance o f wetland plants that b e c o m e established; 3)
h o w long do treatment effects persist; 4) do the treatments increase the abundance o f plant species k n o w n
to be used as food sources b y w a t e r f o w l and marshbirds; and 5) can these techniques be applied at the
scale of whole restoration sites?
METHODS
Small-Scale Study
We conducted the small-scale study on six restoration sites selected f r o m a total o f 21 in our ongoing
study of vegetation responses to restoration (Brown
1995). The study sites are located in Jefferson County,
N e w York (4400'N, 7 5 5 2 ' W ) in the L a k e Ontario
and St. L a w r e n c e River lowlands. At each study site,
Guffin series soils o c c u p y the lowest areas (a fine,
mixed, nonacid, mesic Mollic Epiaquept) grading to
C h a u m o n t series soils in the surrounding uplands (a
fine, mixed, mesic Aeric Epiaqualf).
Each restoration site had a r e m n a n t wetland corridor
located in the drainage ditch constructed to r e m o v e
water from the basin. D r a i n a g e occurred at least 40
years ago at all sites and was probably completed
m u c h longer ago at most sites, but no records are available. The remnant wetland corridors h a v e m a n y m o r e
wetland plant species and species-rich seed banks than
the remaining drained areas that b e c o m e shallow
marsh following restoration o f wetland h y d r o l o g y
(Brown 1995). The original wetland vegetation at
higher elevations was entirely replaced by upland field
plants, as drained areas b e c a m e too d r y to support hydrophytic vegetation. T h e drained areas were used primarily for pasture and forage crops, with v e r y little
cultivation.
The experiment consisted o f two treatments and
three types of controls, all of which w e established in
M a y of 1992 (Table 1A). At e a c h o f the six restoration
sites, plots were established at 15 c m below the restored m a x i m u m water level b y surveying with an
auto-level. In the first treatment, we transplanted soil
blocks by r e m o v i n g all upland soil in 0.25 m 2 plots to
a depth of 15 cm, then transplanting all soil, plant
roots, and rhizomes f r o m an equal soil v o l u m e in the
remnant wetland to the e x p e r i m e n t a l plot. We transplanted the soil in four a p p r o x i m a t e l y equal sections,
which were m o v e d as intact as possible. We replicated
this treatment three times at e a c h o f the 6 wetlands. In
the second treatment, we first sieved the transplanted
soil by passing it through a 1-cm screen to r e m o v e
plant roots and rhizomes. We then transplanted the
sieved soil in the same w a y as the soil blocks. We
replicated this treatment three times at each o f two

426

W E T L A N D S , V o l u m e 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 - 1 5 cm plots from 5
of the 6 sites where a natural drawdown 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
Plots at - 15 cm
Plots at - 3 0 cm
Total Plots
Drawdown Plots
B) Large-Scale Experiment
Treatment
Total Plots

Sieved Soil

Unmanipulated

Litter Removed

24
15

6
0
6
6

18
6
24
15

6
2
8
6

Control

Mowed

Plowed

Soil Trans.

Total

39

30

18

54

141

18

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 disturbance plots had the upland soil to a depth of 15 c m
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 established once at each of the six sites. To address the
influence of hydroperiod on plant responses to these
treatments, we established an additional set o f plots at
30 c m below m a x i m u m water level in two sites, each
with three replicates of transplanted soil blocks and
unmanipulated control plots, and one replicate each o f
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 effects resulted from soil disturbance, litter removal, or
from the transplantation of wetland soil. We analyzed
the effect of disturbing the soil by comparing unmanipulated controls with disturbed soil plots; the effect
o f litter removal by comparing litter removal plots to
unmanipulated controls; the effect of transplanting
wetland soil by comparing the soil-transplant treatment
with unmanipulated controls; and the effect o f including rhizomes in the cores b y comparing the sieved
transplanted soil treatment to the transplanted soil
block treatment.
In October of 1992, after seed germination and seedling 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 perermial plants in the field and identified them when flowers or fruits developed. Taxonomic nomenclature follows
Gleason and Cronquist (1991).

Disturbed
6
2
8
6

We analyzed the data for each plot by calculating a

weighted average index value (Wentworth and Johnson 1986) based on the wetland indicator status o f
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 obligate wetland to obligate upland. The categories reflect
the estimated range o f probabilities o f a species occurring in wetland areas as defined by Cowardin et al.
(1979) as follows: Obligate Wetland (OBL) = > 9 9 % ;
Facultative Wetland (FACW) = 6 7 - 9 9 % ; Facultative
(FAC) = 3 4 - 6 6 % ; Facultative Upland (FACU) = 133%; Obligate Upland (UPL) = < 1 % . M a n y 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 o f its probability range, as follows: O B L
= 1.0; F A C W = 1.67; F A C = 3.0; F A C U = 4.33;
U P L = 5.0. Species with an indicator status o f FAC,
FACW, or O B L 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:

WI=

i=]

(IVl'WIS1)

100

Brown & Bedford, WETLAND

RESTORATION

WITH TRANSPLANTED

SOIL

427

Table 2. Wetland plant species with high value as wildlife food sources that occurred at sites in this study, e = excellent, g
= good, f = fair, p = poor, but important for some species. Compiled from Martin et al. (1951) and Payne (1992).
Genus
Alisma
Bidens
Carex
Comus
Eleocharis
Glyceria
Juncus
Leersia
Lemna
Ludwigia
Polygonum
Polygonum
Potamogeton
Sagittaria
Sparganium

Species
sp.
sp.
sp.
sp.
sp.
striata Lam.
sp.
oryzoides L.
minor L.
palustris L.
hydropiper L.
sp.
sp.
latifolia Willd.
eurycarpum Engelm.

Part Consumed

Waterfowl

seeds
seeds
seeds
fruits
roots
seeds
seeds
seeds, roots
foliage
seeds
seeds
seeds
seeds, roots, stems
seeds, roots
seeds

f
p
f
g
p
f
f
g
g
f
g
e
g
f
f

w h e r e I V = the 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 in that
plot, a n d W I S --- the W e t l a n d I n d i c a t o r Status o f that
species. 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 in
e a c h p l o t w a s u s e d to c a l c u l a t e its i m p o r t a n c e v a l u e in
that plot. W e s u m m e d the p e r c e n t c o v e r o f all s p e c i e s
r e c o r d e d in the 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 the total so i m p o r t a n c e v a l u e s for e a c h
p l o t s u m to 100. 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 to the 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 an
i n d i c a t o r status o f F A C , F A C W , o r O B L a n d the t o t a l
c o v e r o f these w e t l a n d plants.
W e c o m p a r e d the t r a n s p l a n t e d w e t l a n d soil b l o c k s
and undisturbed controls with a two-way ANOVA on
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 t m e n t a n d site
( 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 for
d i f f e r e n c e s in 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 the t o t a l perc e n t c o v e r o f w e t l a n d s p e c i e s in 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 test 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 in e a c h o f the six sites a n d the t w o
f a c t o r s o f p l o t t y p e a n d site. W e u s e d n o r m a l p r o b a -

Marshbirds and
Shorebirds

f
f
e
f

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 uals.
All other comparisons were tested using KruskalW 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 the
s a m p l e sizes 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 1992). W e r e p e a t e d the tests for all
three of the response variables described above (WI,
WSN, and WSC).
Large-Scale Study
W e s e l e c t e d five 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 in t h e
s u m m e r o f 1993 that w e r e s c h e d u l e d for r e s t o r a t i o n
w o r k that fall. E a c h site h a d s o i l s in the s a m e f a m i l y
as the sites in t h e s m a l l s c a l e s t u d y . A t e a c h site, w e
e s t a b l i s h e d soil 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 the e l e v a t i o n s o f t h e site a n d m a r k i n g a n a r e a at l e a s t 1 0 - m
wide and extending across the entire wetland basin
p e r p e n d i c u l a r to the d r a i n a g e c h a n n e l ( b e t w e e n 55 a n d
95 m across). 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 t s for
m o w i n g a n d p l o w i n g at e a c h site a n d a l s o m a r k e d c o n -

Table 3. Results of two-way A N O V A s are shown by year for each response variable for drawdown plots in the small-scale
study. Factors were plot type and site. Plot types compared include only transplanted soil blocks and unmanipulated controls.
R ~ is the value for the model. (* = significant at alpha = 0.05; ** = significant at alpha = 0.01)
Response Variable
Wetland Index
Wetland Species Number
Wetland Species Cover

Year

R~

1992
1993
1992
1993
1992
1993

0.951
0.514
0.816
0.779
0.843
0.706

Site p
p
p
p
p
p
p

=
=
=
<
=
=

0.001"*
0.01"
0.003**
0.001"*
0.001"*
0.014"

Plot type p
p
p
p
p
p
p

<
=
<
<
<
<

0.001"*
0.69
0.001"*
0.001"*
0.001"*
0.001"*

428

W E T L A N D S , V o l u m e 17, No. 3, 1997

a

X
"0

1992

_e 4
0

<
-o

I1

'

.C

0
a

-5

1993

"0

_c 4
01

<
~

t,,,

'~ 1
0

B
S
C
L
D
Transplant Plot Type

Figure 1. Box plots of weighted average wetland index values 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 transplanted with a bulldozer from the remnant wetland in the
drainage ditch in a band that extended upslope to the
edge of the proposed wetland. M o w e d plots had upland vegetation cut but not removed, and plowed plots
had upland soil turned over, both with standard farm
equipment. M o w i n g 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 h y d r o l o g y was restored at all five sites in
the fall o f 1993. During the spring of 1994, we established quadrats at k n o w n elevations at all sites by surveying with an autolevel. Within each treatment area,
we established two replicate transects of 3 plots each
at the elevation o f m a x i m u m water level and at 15 cm
and 30 c m below water level. At one very large site,
we established each treatment twice to test for consistency o f results within a site and installed a set o f
transects in each o f 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

o f three plots. The total numbers o f plots in each treatment are shown in Table lB.
We used the wetland index described above to compare the indicator status o f vegetation in each plot. We
also determined the number of wetland plant species
and the total percent cover o f wetland plants in each
plot. In addition, we determined the number o f species
and the total cover of the plant species k n o w n to be
used as food sources by waterfowl and wetland birds,
as reported from 65 years of field and laboratory research by the U.S. Fish and Wildlife Service on the
food preferences of birds (Table 2).
We compared the wetland index values, number o f
wetland plant species, and total cover a m o n g treatment
types and sites with t w o - w a y A N O V A s using factors
of site and treatment type and repeated each test for
each of the three elevations in each year. We tested the
effects o f elevation on wetland index values using a
two-way A N O V A with factors of treatment type and
plot elevation. We compared total cover o f Typha spp.
among treatment types separately at each elevation using a one-way A N O V A . We used normal probability
plots to test assumptions o f normality for data and residuals. When treatment was a significant effect, we
compared pairwise combinations of treatment types
with "lMkey's HSD procedure. All statistical procedures
were carried out using SYSTAT (Wilkinson 1992).
RESULTS
Small-Scale Study
A natural drawdown of water level occurred during
each summer at five o f the six sites. The experimental
plots at these sites had no standing water for approximately two months during each growing season before reflooding in the fall. All o f 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 ( - 3 0 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 o f the study, and no data
analyses are presented for these plots. Germination o f
seeds in the permanently flooded plots probably was
not triggered without the fluctuating temperature regime and increased oxygen levels that occur when soil
is exposed.

Transplanted Soil Blocks and Unmanipulated Controls. The transplanted wetland soil blocks had sig-

nificantly lower wetland index values than the unmanipulated 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 & Bedford, 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

429

A) Mean N u m b e r of Wetland Plant Species by T r e a t m e n t

7

"6

Block
50 ,

Sieved

Control

Litter

Disturbed

B) Mean Total Cover of Wetland Plants by Treatment

~. so . ~

,,

25
Block

Sieved

Control

Litter

Disturbed

Figure 2. Mean number of wetland plant species (A) and mean total percent cover of wetland plants (B) in drawdown plots
of each type in the small-scale study in each year, with standard deviations. Plot types are as follows: Block = transplanted
soil block; Sieved = transplanted sieved soil; Control = unmanipulated control; Litter = litter removed; and Disturbed =
upland soil disturbed.
wetland species and the amount o f c o v e r of wetland
species in each year (Table 3, Figure 2). Site was a
significant factor in each year, and there were two significant interactions between site and treatment type,
in 1992 for wetland index value (p = 0.016), and in
1993 for wetland species cover (p < 0.001).

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
transplants with only soil and no wetland plant rhiz o m e s . We c o m p a r e d these plots to the unmanipulated
controls and to the transplanted soil blocks within
sites. Transplanting sieved soil lowered the wetland
index value relative to the unmanipulated controls in
the first year, but by the second year, the values were
not significantly different (Table 4A, Figure 1). In the
second year, transplanted soil blocks had significantly
m o r e wetland species than transplanted sieved soil (Table 4A, Figure 2A). In both years, there was significantly more total c o v e r f r o m wetland plant species in
plots with sieved soil than in unmanipulated controls

(Table 4A, Figure 2B). Both the amount o f c o v e r and

the n u m b e r o f species increased at all treatments in the
second year relative to the first year.

Litter Removal and Disturbed Soil Plots Compared to

Unmanipulated Controls. Both litter r e m o v a l and
disturbance o f upland soil affected plant responses in
the first year but not in the second. In both plot types,
wetland index values w e r e significantly lower than in
unmanipulated controls (indicating a greater dominance of wetland plants) but only in the first year (Table 4B, Figure 1). Soil disturbance also increased the
n u m b e r of wetland plant species and the total c o v e r o f
wetland plants relative to unmanipulated controls, but
again only in the first year (Table 4B, Figures 2 A and
B).
Large-Scale Study
In the first year, soil transplant plots at - 1 5 c m had
significantly lower wetland index values than unma-

430

WETLANDS,

Volume

17, N o . 3, 1 9 9 7

Table 4. Results of Kruskal-Wallis o n e - w a y analyses of variance comparing transplant types and control types at d r a w d o w n
plots in the small-scale study for the three response variables: W I = Wetland Index; W S N = Wetland Species N u m b e r ; W S C
= Wetland Species Cover. A) Comparisons between soil transplant types. B) Comparisons b e t w e e n control types. Abbreviations
f o l l o w Table 1. (* = significant at alpha = 0.05; ** = significant at alpha = 0.01)
Year

Comparison

WI

WSN

WSC

A) S i e v e d Transplanted Soil Plots

1992

Trans. B l o c k - - S i e v e d Soil
Sieved Soil--Unmanipulated
Trans. B l o c k s - - S i e v e d Soil
S i e v e d Soil-----Unmanipulated

p
p
p
p

=
=
=
=

0.465
0.004**
0.743
0.949

p
p
p
p

=
=
=
=

0.070
0.127
0.013"
0.099

p
p
p
p

=
=
=
=

0.423
0.004**
0.846
0.048*

B) Manipulated Control Plots

1992
Litter R e m . - - U n m a n i p u l a t e d
Disturbed~Unmanipulated
1993
Litter R e m . - - U n m a n i p u l a t e d
Disturbed--Unmanipulated

p
p
p
p

=
=
=
=

0.032*
0.018"
0.774
0.830

p
p
p
p

=
=
=
=

0.341
0.049*
0.359
0.412

p
p
p
p

=
=
=
=

0.028*
0.028*
0.374
0.200

1993

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 at all 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 terns, with no significant interaction between treatment
t y p e a n d e l e v a t i o n ( F = 1.634, 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 is 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 plants also significantly increased both the number of
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 at all 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 year, n o s i g n i f i c a n t d i f f e r e n c e s in 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 in
p l o t s at - 1 5 c m ( F i g u r e 3). A g a i n , t h e p a t t e r n w a s s i m i l a r at all 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 action between treatment type and elevation (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
plant species was significantly increased by soil transplantation and plowing (Figure 4A). Soil-transplant
plots had significantly more species than control plots
at all 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 at 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 plant plots also had significantly greater cover than
c o n t r o l s at all 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 at all 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
the second growing season, the numbers of species and
t o t a l c o v e r at t h e c o n t r o l p l o t s h a d i n c r e a s e d b u t w e r e
still 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

Table 5. Results of t w o - w a y A N O V A s for the large-scale experiment are shown for each response variable at each elevation
in each year. Factors were treatment type and site, and R: is the value for the model. Treatments included m o w i n g , plowing.
and transplanting wetland soil, and controls were natural, unmanipulated areas. (* = significant at alpha = 0.05; ** = significant
at alpha = 0.01)
Response Variable
Wetland Index

Elev.
Edge
-15em
-30cm

Wetland Species N u m b e r

Edge
-15cm
-30cm

Wetland Species C o v e r

Edge
15cm
-30cm

Year

R2

Site p

Treatment p

1994
1995
1994
1995
1994
1995

0.793
0.751
0.595
0.793
0.605
0.746

p
p
p
p
p
p

==
=
=
<
<

0.014"
0.001"*
0.164
0.002**
0.001"*
0.001"*

p
p
p
p
p
p

<
<
<
=
<
=

0.001"*
0.001"*
0.001"*
0.049*
0.001"*
0.073

1994
1995
1994
1995
1994
1995

0.828
0.730
0.735
0.847
0.815
0.444

p
p
p
p
p
p

<
=
=
=
=
=

0.001"*
0.035*
0.001"*
0.012"
0.002**
0.160

p
p
p
p
p
p

<
<
<
<
<
=

0.001"*
0.001"*
0.001"*
0.001"*
0.001"*
0.034*

1994
1995
1994
1995
1994
1995

0.913
0.869
0.799
0.759
0.701
0.619

p
p
p
p
p
p

=
<
=
<
=
=

0.006**
0.001"*
0.181
0.001"*
0.001"*
0.001"*

p
p
p
p
p
p

<
<
<
=
<
-

0.001"*
0.001"*
0.001"*
0.002**
0.001"*
0.206

B r o w n & Bedford, 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
(Figures 4B and 5B). The number of wetland plant
species decreased in the plowed plots, which were
dominated by cattail.
The responses o f the plant species used by waterfowl and marshbirds as food sources were very similar
to the responses o f wetland vegetation in general. The
soil-transplant plots had significantly greater species
number and total cover than control plots at all elevations in the first year (Figures 6A and 7A). The
m o w e d 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
o f species at the plowed plots had decreased. There
was significantly more cover 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 increased by the plowing treatment (Figure 8). At all
three elevations, the mean total cover o f cattail was
significantly greater in the plowed plots than in each
o f 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 predominantly old-field species in the area that became shallow
marsh following restoration, we hypothesized that
re-establishment o f hydrophytic vegetation would proceed more quickly following transplantation of wetland propagules from the remnant wetland soil. Our
results strongly support this hypothesis and show that
transplantation o f remnant wetland soils can increase
the number and cover o f wetland species in the reflooded areas of restoration sites. The plots with transplanted soil blocks had lower wetland index values,
more wetland species present, and more cover o f wetland 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. Germination of wetland plants, and in particular wetland annuals, occurs most aggressively when standing water is
removed from the soil during a drawdown (Gallinato and

431
a

ab

1994

1995

o) 3

<[2
>

C M P ST
TreatmentType
I

Figure 3. Box plots of weighted average wetland index values for plots at - 1 5 c m in each large-scale experiment plot
type in 1994 and 1995. Plot types are as follows: C = unmanipulated control; M = mowed; P = plowed; ST = wetland 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 establishment 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 eventually become established at lower elevations as well, but
the study period was too short to observe this aspect of
plant community 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 transplantation of wetland soil. By the second year, however, manipulated upland plots without soil amendments showed no improvement in any of the response
variables relative to the unmanipulated controls. After
two years of flooding, e n o u g h wetland plant establish-

432

W E T L A N D S , V o l u m e 17, No. 3, 1997

14
12

A) Mean Number of Wetland Plant Species in 1994

a

aab

bc c

b c

[ ] Control
13Mowed
E] Plowed

[] Soil Transplant

-30

14
12
10

-15

Edge

B) Mean Number of Wetland Plant Species in 1995

a

a a

aabb

ili_=
-30

aab

--

II Control
[ ] Mowed
[] Plowed
I~ Soil Transplant

-15

Edge

Figure 4. Mean number of wetland plant species in 1994 (A) and 1995 (B) in plot types in the large-scale experiment, with
standard deviations. Treatments within each elevation that do not share a common letter are significantly different at alpha =
0.05.
ment had occurred in the unmanipulated controls to
m a k e them similar to the litter-removal and soil-disturbance plots. Thus, it seems that while t e m p o r a r y
effects can be produced by removing litter or disturbing the upland soil, only the addition of soil f r o m the
remnant wetland has longer term effects on the wetland plant c o m m u n i t y that develops. Transplantation
of remnant wetland soil seems to be the m o s t effective
technique for increasing the re-establishment of wetland plants relative to natural recolonization.
There were no differences between plots with transplanted soil and plots with transplanted soil and rhizomes
in the first year, with both types showing significant increases in number and total cover of wetland plant species compared to unmanipulated controls. These results
suggest that, during the first growing season following
transplantation, there is no practical difference in the
re-establishment of wetland plants that results from these
two methods. However, in the second year, the plots with
transplanted rhizomes had significantly greater numbers
of wetland species than those with only sieved soil. This

suggests that over a longer term, rhizomatous species can

become established and increase the species density in
plots where they are transplanted. Long-term storage of
soil before transplantation at restoration sites could reduce the number of viable rhizomes and therefore reduce
the number of successfully established species. This m a y
be significant since rhizomes are associated with perennial emergents that are important for restoration of avifauna food and cover (Martin et al. 1951, Payne 1992).
The results f r o m the small-scale experiment show
that the three response variables o f n u m b e r of wetland
plant species, amount of cover, and wetland index all
reflect different characteristics of the restored plant
community. In the second year of the study, wetland
plants at transplanted soil blocks had greater c o v e r
than at unmanipulated controls, but the wetland index
values for the two types o f plots did not differ. This
was true even though the wetland index values were
calculated using the relative c o v e r o f each species.
Within two growing seasons after wetland hydrology
was restored, only plant species that could tolerate sat-

B r o w n & Bedford, 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
A ) Mean Total Cover of W e t l a n d Plants in 1994

300
250

a ab bc c

200
"E

Control
IR Mowed

150

[ ] Plowed
[] Soil Transplant

n
I--

433

100
50

-30

>

Edge

B) Mean Total Cover of W e t l a n d Plants in 1995

300
250

-15

a abab

a ab b

200

Control
Mowed

150

rlPIowed

O tii/

[ ] Soil Transplant

"~ 100
50

-30

-15

Edge

Figure 5. Mean total percent cover of wetland plants in the large-scale experiment in 1994 (A) and 1995 (B), with plot
layout following Figure 4.

urated soils persisted, so the index values of the vegetation present were uniformly low across all treatment
types. At the same time, the transplanted soil blocks
had significantly more cover o f wetland plants and
m o r e plant species. The index values reflected the
changes in vegetation that occurred in the first growing
season following restoration of hydrology. However,
the index value did not reflect the differences in the
wetland plant c o m m u n i t y that persisted after the majority o f surviving plants were wetland species. Analysis of restored plant communities involving comparison with reference wetlands should include measurement of species richness and cover because index values 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 in
c o m m u n i t y structure after the initial readjustment to
wetland conditions.
Large-Scale Study
The large-scale study confirmed that the increases
in wetland plant responses observed in the small-scale

study could be achieved at the scale o f whole sites.

The large-scale results showed that soil transplantation
can increase the n u m b e r and the c o v e r of wetland plant
species at restored wetlands and that the technique can
be applied at the scale of whole sites during initial
restoration work. I m p r o v e m e n t in avifauna habitat requires increases in the n u m b e r and c o v e r o f wetland
plants that are used as f o o d sources. The transplant
treatments significantly i m p r o v e d both the n u m b e r of
species and the amount o f c o v e r of food plants. This
treatment could significantly i m p r o v e the habitat value
o f restored sites for avifauna, particularly in the first
few years. Siegley et al. (1988) found similar results
for marshes created using dredge spoils on L a k e Erie
that contained wetland plant seeds, which showed increases in wildlife food plant d e v e l o p m e n t when seedrich spoils were used.
However, we do not k n o w h o w long the effect of
soil transplants will persist, and short-term establishm e n t patterns m a y not predict long-term plant communities (Weiher et al. 1996). It seems likely f r o m the

434

W E T L A N D S , V o l u m e 17, No. 3, 1997

A) Mean N u m b e r of Wildlife Food Plants in 1994
a

a abab

a abab

III Control
1
[] Mowed
/
r-I Plowed
/
[] Soil Transplant j

T
-30

Edge

-15

B) Mean N u m b e r of Wildlife Food Plants in 1995

a

a ab b

[]
[]
[]

-30

-15

Control
Mowed
Plowed
Soil Transplant i

Edge

Figure 6. Mean number of wildlife food plant species in the large-scale experiment in 1994 (A) and 1995 (B), with plot
layout following Figure 4.

results at other restored wetlands in our study that the

untreated areas will develop similar species richness
and cover of herbaceous wetland plants over approximately three years (Brown 1995). I f the i m p r o v e m e n t
f r o m soil transplants does not persist longer than the
first few growing seasons, then the technique would
be unnecessary as a general practice.
The most important use o f this technique m a y be to
control cattail in areas where soil has been disturbed
during construction. Soil disturbance in the plowed
treatments had the predicted effect of increasing colonization by seedlings of Typha spp., and we measured
significantly lower numbers and cover of wildlife food
plants in these areas. This has been a significant problem at other restoration sites in our study, particularly
in the area where dike construction disturbs the soil
(Brown 1995). Monotypic stands of cattail reduce vegetation diversity and the ratio of open water to vegetation, and, therefore, the habitat value of the site (Weller 1975, Kaminski and Prince 1981, Murkin et al.
1982). R e d u c t i o n o f cattail t h r o u g h the u s e o f

coarse-textured substrates, as suggested by Weiher et

al. (199"6), m a y be effective at created wetlands but
would be cost-prohibitive for large-scale restoration
projects. Transplanting r e m n a n t wetland soil into the
area disturbed during dike construction should substantially i m p r o v e the establishment of other m o r e
valuable wetland plant species in these areas and limit
populations of cattail. Vivian-Smith and Handel (1996)
also r e c o m m e n d the use of imported soils to i m p r o v e
wetland restoration sites. E v e n though this technique
increases the initial cost of construction, it m a y prevent
the need for costly intensive m a n a g e m e n t techniques
to control cattail and can be accomplished while construction equipment is on site.
The effect of transplanted wetland soils on plant recruitment in a restored wetland will v a r y depending
on the composition of the seed bank. We studied the
seed bank in the remnant wetland soils before undertaking this experiment and found that they contained
a variety of wetland plant species useful for habitat
i m p r o v e m e n t (Brown 1995). In addition, the seed b a n k

B r o w n & Bedford, 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

435

A) M e a n Total C o v e r of Wildlife F o o d Plants in 1 9 9 4

a abab

a abab

a abab

~> 1
o
o

II
LT I

-30
175
150

[] Control
[]Mowed
I'llPIowed
[] Soil Transplant

T
Edge

-15

B) M e a n Total C o v e r of Wildlife F o o d Plants in 1995

aacbc

a a b c

aabb

125
100

g.

ol

=Mowed

t"llpIowed
[] Soil Transplant

-30

-15

Edge

Figure 7. Mean total percent cover of wildlife food plants in the large-scale experiment in 1994 (A) and 1995 (B), with plot
layout following Figure 4.
lacked undesirable invasive species such as purple
loosestrife (Lythrum salicaria L.). Transplanting remnant wetland soil is prudent only when the composition of the seed b a n k is k n o w n and is consistent with

200
180

m a n a g e m e n t goals (Pederson and van der Valk 1984,

van der Valk et al. 1992). F o r example, transplanting
remnant wetland soil f r o m areas dominated by cattail,
which are likely to have depauperate seed banks and

M e a n Total C o v e r of
a

Typha spp.
a

in 1995
b

160
o~ 140

~ 120
g-5

I---

100

8o
60
4O
20
0
-30

Figure 8.

-15

IBControl
I
a]Mowed
/
131Plowed
]
[] Soil Transplant I

Edge

Mean total percent cover of Typha spp. in the large-scale experiment in 1995, with plot layout following Figure 4.

436

WETLANDS,

large numbers of cattail seeds, would probably not inc 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 as e f f e c t i v e l y .

T h e w a t e r l e v e l is 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
seed germination and, therefore, plant establishment in
a wetland (Keddy and Ellis 1985, Moore and Keddy
1988). In our study, establishment of wetland plants
decreased steadily with increasing depth relative to
maximum high water. We observed little establishment
at water depths greater than 45 cm even though the
treatments extended across the entire wetland basin,
This suggests that transplantation of wetland soil
should be concentrated at shallower elevations near the
proposed high water level of restored sites.
Relocation of wetland soil should be considered as
a potentially useful restoration technique whenever
t h e r e is 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 site. S e e d
bank assays should be conducted to ensure that the
s p e c i e s c o m p o s i t i o n is 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
restoration project. The use of remnant seed banks provides a relatively low-cost source of wetland plant
propagules that can significantly increase the restoration of plant communities at drained wetlands. The
t e c h n i q u e is 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 lishment of monotypic cattail stands in areas where
soil has been disturbed by dike construction.
ACKNOWLEDGMENTS
This research was supported by Region 5 of the U.S.
Fish and Wildlife Service with additional financial
support from Ducks Unlimited, Inc., the National Fish
and Wildlife Foundation, the Edna Bailey Sussman
Fund, and Cornell University. Our field assistants, including Bill Yamell, Metta McGarvey, and Christian
Ottke, worked tirelessly, cheerfully, and often independently and contributed immeasurably to the project. Karen Poiani provided advice and moral support
throughout the project and provided detailed comments on the manuscript. The manuscript also benefited from the comments and suggestions of two anonymous reviewers.
LITERATURE

CITED

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control by burning and mowing. Journal of Aquatic Plant Management 28:32-36.
Beule, J. D. 1979. Control and management of cattails in southeastern Wisconsin wetlands. Department of Natural Resources,
Madison, WI, USA. Technical Bulletin No. 112
Brown, S. C. 1995. Wetland restoration: Factors controlling plant
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Comes, R. D. and A. D. Kelly. 1989. Control of common cattail
with postemergence herbicides. Journal of Aquatic Plant Management 27:20-23.
Cowardin, L. M., V. Carter. E C. Golet, and E. T. Laroe. 1979.
Classification of wetlands and deepwater habitats of the United

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FWS/OBS-79/31.
Dunn, W. J. and G. R. Best. 1984. Enhancing ecological succession.
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Sedimentology, and Reclamation. Office of Engineering Services,
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Eicher, A. L. 1988. Soil-vegetation correlations in wetlands and adjacent uplands of the San-Francisco Bay estuary. California. U.S.
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6:102-112.
Gallinato, M, I. and A. G. van der Valk. 1986. Seed germination
traits of annuals and emergents recruited during drawdowns in the
delta marsh, Manitoba, Canada. Aquatic Botany 26:89-102.
Gleason, H. A. and A. Cronquist. 1991. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. New York
Botanical Garden, Bronx, New York, NY, USA.
Harris, S. W. and W. H. Marshall, 1963. Ecology of water level
manipulations on a northern marsh. Ecology 44:331-343.
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