Journal of Applied

Assessing grassland restoration success: relative roles of

Blackwell Publishing Ltd

Ecology 2006
seed additions and native ungulate activities
LEANNE M. MARTIN and BRIAN J. WILSEY
Iowa State University, Department of Ecology, Evolution and Organismal Biology, Ames, IA 50011, USA

Summary
1. Grassland restorations often lack rare forb and grass species that are found in intact
grasslands. The possible reasons for low diversity include seed limitation, microsite
limitation and a combination of both. Native ungulates may create microsites for
seedling establishment in tallgrass prairie restorations by grazing dominant species or
through trampling activities, but this has never been tested in developing prairies.
2. We experimentally tested for seed and microsite limitation in the largest tallgrass
prairie restoration in the USA by adding rare forb and grass seeds in two trials inside
and outside native ungulate exclosures. We measured seedling emergence because this
stage is crucial in recruiting species into a community. We also measured light, water
and standing crop biomass to test whether resource availability could help to explain
seedling emergence rates.
3. Ungulates increased light availability for each sampling time and also increased
above-ground net primary productivity (ANPP) during summer.
4. Seedling emergence of rare prairie forbs and grasses was consistently greater when
we added seeds.
5. Seedling emergence was conditionally greater with a combination of seed additions
and grazing, but grazing alone was unable to increase emergence.
6. When ungulates increased seedling enhancement, the mechanism was partially
associated with increased water and light availability.
7. Exotic and cosmopolitan weed seedling emergence was not affected by grazing.
8. Synthesis and applications. These results suggest that tallgrass prairie restorations
are primarily seed limited and that grazing alone may not be able to increase seedling
emergence of rare species without the addition of seeds. Therefore, adding seeds to
grassland restorations may increase seedling emergence of rare species, and mimicking
effects of grazing may increase emergence when seeds are added.
Key-words: Bos bison, Cervus elaphus, diversity, grazing, Iowa, net primary productivity,
seedling emergence, tallgrass prairie
Journal of Applied Ecology (2006)
doi: 10.1111/j.1365-2664.2006.01211.x

establish reproducing populations, and then popula-

Introduction
Ecosystem restoration is becoming a more common
way to increase native species habitat. Typically, resto-
rations are attempted by adding seeds from nearby
remnants to a previously converted ecosystem (Sluis
2002; Polley, Derner & Wilsey 2005). Seedlings of
multiple species are expected to emerge, survive and
© 2006 The Authors.
Correspondence: Leanne M. Martin, University of
Journal compilation Nebraska at Omaha, Department of Biology, 6001 Dodge
© 2006 British St, Omaha, NE 68182, USA (fax 402 554 3532; e-mail
Ecological Society lmmartin@mail.unomaha.edu).
tions are expected to assemble into a community
similar to the original system. The seedling emergence
stage is important in this process because it funnels
individuals into the system. Contrary to expectations,
restored ecosystems often have lower plant species
richness and diversity than their unaltered counterparts
(Galatowitsch & van der Valk 1996; Martin, Moloney
& Wilsey 2005; Polley, Derner & Wilsey 2005) and
species richness has been observed to decline over time
instead of increase as expected (Sluis 2002).
Typically, low species diversity is attributed to either
(i) seed limitation or (ii) seedling microsite limitation
2 (Eriksson & Ehrlen 1992; Zobel et al. 2000; Foster
L. M. Martin & et al. 2004; Henry et al. 2004). The seed limitation
B. J. Wilsey hypothesis suggests that plant community richness and
diversity are limited by the species pool (Gough, Grace
& Taylor 1994). Seed additions have increased species
richness and diversity of some native plant commun-
ities and agriculturally improved grasslands (Pywell
et al. 2002; Smith et al. 2002; but see Wilsey & Polley
2003). If restorations are seed limited, then adding
seeds of a large number of species should increase
diversity and recruitment of rare species even in sys-
tems with high dominance. Alternatively, the microsite
hypothesis suggests that one or a few strongly dominant
species suppress seedlings (Howe 2000; Sluis 2002;
Camill et al. 2004). In this scenario, seeds or prop-
agules are not limiting but seedlings fail to establish
reproducing populations. Dominant grass patches are
usually larger in restorations than in intact grassland
(Derner et al. 2004). Dominance by C4 grasses, which
can occur as soon as 3 years after establishment, can be
especially high in the nutrient-rich environments that
characterize most restorations (Baer et al. 2002; Baer
et al. 2004; Camill et al. 2004). Large grass canopies
and abundant litter can reduce light and water avail-
ability, which are crucial to seedling survival (Fahnestock
& Knapp 1993; Haugland & Froud-Williams 1999;
Xiong & Nilsson 1999). Common management practices,
such as frequent spring burning and grazing exclusion,
could exacerbate this problem (Collins et al. 1998; Howe
2000). Thus, anything that reduces grass dominance
should alleviate competition with rare species and
should increase seedling establishment and diversity
(Foster & Gross 1997).
According to the intermediate disturbance and
grazing optimization hypotheses, intermediate levels
of grazing should produce the highest levels of species
diversity, but also the highest levels of net primary
productivity (NPP) in an ecosystem (Grime 1973;
Connell 1978). The question of whether native ungulate
grazing can increase seedling emergence and diversity
is becoming more relevant because grazers such as
bison Bos bison L. and wapiti Cervus elaphus L. are
increasingly being reintroduced (Knapp et al. 1999;
Larkin et al. 2004). Management strategies, such as
mowing, aimed at decreasing the biomass of dominant
species have shown increased seedling survival in
some experimental and pasture plantings (Burke &
Grime 1996; Hutchings & Booth 1996; Lawson, Ford &
Mitchley 2004). Moderate grazing by native ungulates,
a common grassland disturbance, could have positive
impacts on plant species diversity in intact grasslands
by reducing dominant grasses and increasing light
availability (Hartnett, Hickman & Walter 1996; Collins
© 2006 The Authors. et al. 1998; Knapp et al. 1999). Moderate grazing
Journal compilation
would therefore be expected to produce non-linear
© 2006 British
Ecological Society,
effects on diversity in restorations, with higher levels of
Journal of Applied diversity at intermediate grazing intensities (Smith
Ecology et al. 2000). However, this may be restricted to intact
systems, where there is a propagule source available for
recruitment into the community (Hartnett, Hickman &
Walter 1996; Collins et al. 1998). Furthermore, because
intermediate levels of grazing can also produce the
highest levels of NPP (McNaughton 1979; Dyer,
Turner & Seastedt 1993), grazing may lead to increased
resource uptake by plants. Productivity is already
high in grassland restorations, often higher than in
comparable remnants (Baer et al. 2002; Camill et al.
2004; Martin, Moloney & Wilsey 2005). If inter-
mediate grazing increases production of dominant species
above and beyond what is already high, then interme-
diate grazing in restorations, unlike in intact grasslands,
may actually lessen positive effects on seedling emergence
and diversity. Thus increased productivity in restorations
because of moderate grazing might nullify potentially
positive effects of grazing on microsite availability.
Finally, seed limitation and low seedling emergence
because of grass dominance may interact to limit diver-
sity in grassland restorations. A combination of adding
seeds and increasing microsite availability may be
necessary to favour seedling emergence (Burke &
Grime 1996; Turnbull, Crawley & Rees 2000; Foster &
Dickson 2004).
Our objectives were to determine: (i) if native ungu-
lates increase availability of resources crucial to seed-
ling emergence and (ii) whether seed additions, native
ungulate grazing or a combination of both enhance
native prairie seedling emergence while having little or
no effect on non-native and cosmopolitan weeds in tall-
grass prairie restorations. Our focus was on seedling
emergence, a key stage in the establishment of grass-
land plants. Whether seedlings can establish viable
populations is a longer term question that will not be
considered here.
Materials and methods
 
The objective of the Neal Smith National Wildlife
Refuge (NS) prairie project is to restore a large tallgrass
prairie ecosystem using locally collected seeds combined
with prescribed fire and grazing by native ungulates.
The restoration is located on the Walnut Creek water-
shed in Jasper County, Iowa, USA (41°33′N, 93°17′W).
The refuge currently spans 2104 ha, approximately
1200 ha of which have been seeded with tallgrass
prairie species, beginning in 1992 and continuing to the
present day. Grazing ungulates (B. bison and C. elaphus)
were introduced to a 303-ha enclosure in 1996 and
1998, respectively, which is where our study took place.
Approximately 35 B. bison and 15 C. elaphus occupied
the area during our study. Land use prior to prairie
seeding included corn Zea mays L. and soybean Glycine
max (L.) Merr rotations and a few scattered pastures.
There were 20 different plantings in this area (mean of
approximately 14 ha each) and each planting was seeded
with separate bulk seed mixes collected from local prairie
remnants. Management practices after planting included
3 yearly spring burning during the early years followed
Native ungulates by 2-year burn rotations, which is a common practice
and seed additions for beginning restorations (Packard & Mutel 1997;
in restorations Copeland, Sluis & Howe 2002). Mowing was done
when necessary to control weedy and invasive species
(P. Drobney, personal communication). Our plots were
not burned or mowed in 2003 or 2004, the years of
sampling.
Historically, precipitation at the site has a unimodal
distribution and peaks in May and/or June, with an
average of approximately 880 mm year−1. Weather in 2003
was much warmer and drier than during 2004. Between
May and August, the peak growing months, tempera-
tures and monthly precipitation averaged 21·7 °C and
13·3 mm in 2003 and 19·7 °C and 143·8 mm in 2004.
To standardize our sampling, we randomly selected
eight plantings within the enclosure that were seeded
between 1994 and 1996 on formerly cropped areas.
Four plantings north of a dirt road included 6·7 kg ha−1
of Elymus canadensis L. in the seed mixture as a puta-
tive cover crop, whereas four plantings south of the
road did not (for effects of the cover crop see Martin,
Moloney & Wilsey 2005).
 
A randomized complete block split-plot design with
unequal replication was used, with grazing or exclosures
applied to main plots and seed addition treatments
(described below) applied to subplots. Two 6 × 8-m
grazed plots were established 5 m away on either side of
a permanent 6 × 8-m permanent exclosure in June 2003
in each of the eight plantings (blocks). Two grazed
plots were sampled per planting because of increased
heterogeneity with grazing (Knapp et al. 1999). By
request of the refuge staff, exclosures were kept out of
view of visitors where possible and this precluded
completely random locations.
   
Biomass and above-ground net primary productivity
(ANPP) (general indicators of resource uptake and
competition intensity) were estimated to compare
grazed and exclosed plots (Baer et al. 2004). Above-
ground biomass was clipped to 2 cm in a 40 × 100-cm
quadrat randomly placed in each exclosure (ng) and
grazed plot (gr), and surface litter was collected in June
and August 2003 and in March, June and August 2004.
Biomass was sorted into live and dead components,
and live material was sorted by species, dried for 48 h
at 65 °C, and weighed. Subsequently, estimates were
made for the following biomass components: propor-
© 2006 The Authors. tion of exotic biomass (exotic/total), proportion of
Journal compilation
total grass biomass (grass/total) and C4 grass biomass
© 2006 British
(C4/total), and combined litter and standing dead bio-
Ecological Society,
Journal of Applied mass. Each of these different components of biomass
Ecology could suppress seedling emergence and species richness
in grassland restorations (Howe 2000; Camill et al.
2004). Plant species were designated as native or exotic
based on Eilers & Roosa (1994).
We estimated grazing intensity (GI) and used poly-
nomial regressions to determine if GI was quadratic-
ally related to response variables (McNaughton 1979,
1985). Simply comparing grazed and ungrazed plots
can be misleading in cases where grazing is non-linearly
related to response variables (Grime 1973; Connell
1978; McNaughton 1979). Above-ground NPP and
GI were estimated for grazed plots (n = 16) using the
moveable exclosure approach (McNaughton 1985;
McNaughton, Milchunas & Frank 1996) during three
periods: June–August 2003 and March–June and
June–August 2004. One 3 × 4-m temporary exclosure
was established at each site in March 2004 and was
moved in June 2004 to measure consumption and GI
(McNaughton 1985). Above-ground biomass from the
centre of each temporary exclosure was collected using
the same quadrat size as explained above. Biomass from
the permanent exclosure was used to estimate consump-
tion during June–August 2003 (i.e. the first growing
season). Consumption (C) was estimated as (ng – g)/
time, where ng was biomass inside and g was biomass
outside temporary exclosures at the end of the period,
and time was the number of days exclosures were in
place (McNaughton 1985; Wilsey et al. 2002). Above-
ground NPP (g m−2 day−1) was calculated as a positive
biomass increment + consumption for each time period.
Grazing intensity was calculated as GI = C/NPP
(McNaughton 1985; Wilsey et al. 2002), with GI set to
0 if consumption estimates were negative.
Environmental variables were measured to determine
whether grazing was creating microsites favourable for
seedling emergence. Soil moisture and percentage light
at the soil surface were measured monthly from July to
September 2003 and from May to October 2004 (soil
moisture was not measured in July–August 2003 because
of equipment failure) in each plot using a Moisture
Point® MP-917 Time Domain Reflectometry system
(30 cm rods; Environmental Sensors, Victoria, Canada)
and a 1-m Decagon® AccuPar Ceptometer (Decagon
Devices Inc., Pullman, WA). Sampling points were
randomly located, and two measures of incident light
were taken during each sampling time.
Species diversity was calculated from biomass to
determine if grazing exclusion affected diversity.
Diversity was calculated at the quadrat scale for each
grazed and exclosed plot. Diversity was quantified
2
with Simpson’s diversity (1/D), where D = 1/ ∑ pi and
pi = relative biomass of each species i, and was then
decomposed into species richness (S) and evenness
(1/D/S) to determine if each component of diversity
differed (Buzas & Hayek 1996; Smith & Wilson 1996;
Martin, Moloney & Wilsey 2005).
  
Two seed additions of rare native prairie forbs and
grasses were made to separate, randomly located 1-m2
4 subplots within each plot using two different methods.
L. M. Martin & These were compared to one control subplot (no seed
B. J. Wilsey addition) within each main plot. Therefore, three
subplots were located in each main plot (exclosure or
grazed plot), with a total of nine subplots in each block.
The first seed addition treatment consisted of adding
seeds of 10 species collected by hand from local rem-
nants in June 2003. The second treatment consisted of
adding seeds of 25 species from a local seed company
(Allendan Seed Co., Winterset, IA, USA) in April 2004
in a second set of subplots. More species were used in the
second trial than the first because seeds were more readily
available from the seed company, and we wanted to
mimic the high number of species found at the
neighbourhood scale in remnants (Martin, Moloney &
Wilsey 2005). Seeds were added with equal relative
abundances at a rate of 19 700 seeds m−2 for both trials
(1970 and 788 per species for addition experiments 1
and 2, respectively). Seed numbers were based on
number of seeds in a typical seed rain rate found by
Rabinowitz & Rapp (1980) in a Missouri tallgrass
prairie. Seed viability was not tested with seeds collected
from remnants, but all but one species readily germi-
nated in greenhouse pots grown for seedling reference
samples. Mean seed viability for seeds obtained from
the seed company was 81% (range 49–96%). Seeds
were hand-scattered in each 1-m2 subplot and existing
vegetation and litter were shaken to aid seeds in
reaching the soil surface. Species added in the first
experiment were Bouteloua curtipendula (Michx.)
Torrey, Sporoblus asper (Michx.) Kunth, Solidago
speciosa Nutt., Pycnanthemum virginianum (L.) Dur. &
Jackson, Dalea purpurea Vent., Chamaecrista fasciculata
(Michx.) Greene, Amorpha canescens Pursh, Lespedeza
capitata Michx., Monarda fistulosa L. and Eryngium
yuccifolium (Michx.) (Eilers & Roosa 1994). Species
added in the second experiment included all those
added in the first experiment, plus Potentilla arguta
Pursh, Silphium laciniatum L., Echinacea pallida Nutt.,
Ratibida pinnata (Vent.) Barnh., Artemesia ludoviciana
Nutt., Liatris pycnostachya (Michx.), Verbena stricta
Vent., Helianthus rigidus (Cass.) Desf., Gentiana
andrewsii Griseb., Tradescantia bracteata Small, Viola
pedatifida G. Don, Anemone cylindrica Gray, Phlox
pilosa L., Schizachyrium scoparium (Michx.) Nash and
Solidago rigida L. (Eilers & Roosa 1994).
All forb seedlings, including species added from the
mix as well as volunteers, were identified to species and
counted in a randomly placed 20 × 50-cm quadrat
within each subplot, to estimate seedling emergence.
Volunteers were included because some added species
were already in the seed bank and therefore could not
be distinguished from experimentally sown seedlings.
© 2006 The Authors. Exotic and cosmopolitan weed seedlings were counted
Journal compilation
to test concerns about disturbance from native ungu-
© 2006 British
Ecological Society,
lates increasing weeds in grasslands (Smith & Knapp
Journal of Applied 1999). Grass seedlings were only counted if the species
Ecology was added. Seedlings were counted if they were up
to 7·5 cm tall or up to any height if they were annuals,
and were counted once per month during the grow-
ing season, beginning the month after seeds were
added.
 
Randomized block split-plot s were used, with
planting as a random block term. All grazing effects were
tested with the main plot error term (planting × grazed),
and seed and seed–grazed interactions were tested with
the subplot error term. Repeated-measures 
was used to compare grazed (n = 16) and exclosed plots
(n = 8) for existing vegetation and resource variables,
with time 0 data (measurements taken before exclos-
ures were constructed) as a covariate (except for NPP,
for which time 0 data could not be calculated). We
dropped the covariate from each model if it was not
significant (P > 0·05). Variables were logarithmically
transformed (biomass, standing dead and litter), square-
root transformed (proportion of exotic biomass) or
arcsin square-root transformed (proportion of C4 and
grass) to improve normality when necessary. All
analyses were done with   in SAS (Littell,
Stroup & Freund 2002).
The first and second seed additions were analysed
separately because they had different numbers of spe-
cies, addition dates and weather conditions. A seedling
enhancement effect, ln[(added seedlings + 1)/(control
seedlings + 1)], was calculated to quantify the number
of seedlings that did not emerge from the existing seed
bank but emerged from added seeds. This derived vari-
able eliminated non-normality in data as a result of
having many zeroes in control subplots. To test if seed
additions increased seedling numbers above those of
controls, seedling enhancement effects were tested against
0 with a t-test. Grazing effects on seedling enhancement
over time were analysed with repeated-measures 
of corresponding data. Exotic seedling and seedling
diversity variables were analysed with repeated-measures
(means for 2003 and 2004)  for the first seed addi-
tion, and with regular  for the second addition.
Data were averaged across months because raw data
had too many zeroes to analyse each sampling time.
Seedling Simpson’s diversity (1/D), species richness (S)
and evenness (1/D/S ) were calculated in each subplot
to determine if seed additions or grazing improved
seedling diversity.
We used polynomial regression to test for quadratic
and linear relationships between mean GI (grazed plots
only) and response variables. Mean GI was calculated
by averaging GI across time because of non-normally
distributed data. We used path analysis to test for direct
and indirect associations of grazing on seedling
enhancement. A direct pathway was tested of GI on
biomass, biomass on light and water availability, and
light and water on seedling enhancement. An indirect
path was tested of GI on NPP, and NPP on seedling
enhancement. This indirect pathway could be signifi-
cant if increased NPP had additional effects on seedlings
 Table 1.  results (F-values) for comparisons between grazed plots and exclosures in a tallgrass prairie restoration. Time 0 data were included as a covariate, but were removed if not significant (P > 0·05;
5 because of non-light and water effects, such as increased

Water

5·6*
0·0

1·9
0·9

1·4
Native ungulates nutrient uptake of vegetation.
and seed additions
in restorations

Alpha S
Results

8·0*
1·9

0·8
1·0

1·5
   

Proportion
Only two response variables differed significantly
between grazed and exclosed plots and another was

grass

6·7*
marginally significant (Tables 1 and 2). Above-ground

0·8

0·3
0·1

2·3
NPP m−2 was 1·2, 1·1 and 8·0 times as large in grazed
plots depending on time period, and the difference was

Proportion
only significant for June–August 2004 (Table 1 and

exotic S

20·6**

2·7†
Fig. 1a). Light availability at the soil surface was 1·7

2·4
2·7

0·7
times as great in grazed plots, and this was fairly
consistent across time periods (Table 1 and Fig. 1b).

d.f.
Combined standing dead and litter biomass m−2 was

2
1
14
2

2
28
marginally significantly lower in grazed plots (Tables 1
and 2).

Light

8·8**
6·7*
NS

1·4
No response variables were quadratically related to
mean GI (grazing intensity) (F1,15 between 0·03 and

1
15
6

6
132
d.f.
3·07, P between 0·10 and 0·88). Mean GI was highest in
June–August 2004, when 68% of NPP was consumed
(range 0–100%, SE 6), followed by 49% during June–

Alpha E
August 2003 (range 0–100%, SE 8). It was much

NS
0·1

0·2

0·4
lower during spring 2004 (mean 14%, range 0–37%,
SE 3). Biomass m−2 was negatively related to mean GI

Alpha
denoted by NS) Significance is indicated by †P between 0·06 and 0·1, *P between 0·02 and 0·05 and **P ≤ 0·01

1/D

NS
2·4

1·0

0·1
Proportion
C4 grass

NS
1·8

2·1

0·7
exotic biomass
Proportion

6·3**
NS
0·6

0·8
standing dead
Litter and

3·7†
NS

0·9

1·6
ANPP

9·4**

7·2**
4·3*
NS
Biomass

NS
2·4

1·0

0·1
d.f.

1
15
2

2
44

© 2006 The Authors.

Journal compilation
Time × grazed

© 2006 British
Time × time 0

Error (time)

Ecological Society,
Journal of Applied Fig. 1. Grazing (n = 16) or exclosure from grazing (n = 8)
Grazed
Time 0
Source

Error
Time

Ecology differences for (a) ANPP (P < 0·01) and (b) percentage light
availability at soil surface (P < 0·01). Vertical bars are ± 1 SE.
6

October 2004
August 2004

6·12 (0·17)
6·58 (0·14)
0·22 (0·07)
0·36 (0·04)
1·08 (0·08)
1·01 (0·11)
11·02 (1·10)
0·24 (0·03)
2·76 (0·44)

28·23 (1·40)
L. M. Martin &
B. J. Wilsey

September 2004

34·69 (1·40)
5·97 (0·17)
6·58 (0·14)
0·31 (0·07)
0·31 (0·04)
1·19 (0·08)
0·99 (0·11)
9·86 (1·10)
0·25 (0·03)
2·46 (0·44)
June 2004

33·82 (1·40)
July 2004

35·75 (1·40)
June 2004
Exclosed (n = 8)

August 2003

6·13 (0·17)
6·36 (0·14)
0·17 (0·07)
0·30 (0·04)
1·22 (0·08)
1·13 (0·11)
9·44 (1·10)
0·23 (0·03)
2·27 (0·44)

38·41 (1·40)
September 2003 May 2004
Exclosed (n = 8)

39·98 (1·40)
Table 2. Mean (SE) of response variables measured to test effects of native ungulate grazing in a tallgrass prairie restoration

August 2004

5·93 (0·14)
6·39 (0·14)
0·32 (0·06)
0·36 (0·03)
1·17 (0·06)
0·96 (0·08)
10·86 (0·82)
0·26 (0·02)
2·63 (0·33)

Fig. 2. Relationships between (a) biomass and (b) evenness

and grazing intensity (n = 16).
September 2004 October 2004

26·66 (1·04)

(F1,15 = 6·02, P = 0·03, r = −0·56, slope = −3·49; Fig. 2a),

as expected. Species evenness at the 0·4 m −2 scale
was positively related to mean GI (F1,15 = 5·61, P = 0·03,
r = 0·54, slope = 0·89; Fig. 2b).
5·82 (0·14)
6·09 (0·14)
0·33 (0·06)
0·37 (0·03)
1·01 (0·06)
0·83 (0·08)
12·63 (0·82)
0·24 (0·02)
2·89 (0·33)

35·10 (1·04)
June 2004

- 

Native species seedling emergence

34·84 (1·04)
July 2004

Seed additions increased the number of native species

seedlings 0·1 m−2 in both seed addition experiments
Grazed (n = 16)

(Table 3 and Fig. 3). Adding seeds increased seedling

August 2003

5·87 (0·14)
6·28 (0·14)
0·17 (0·06)
0·35 (0·03)
1·13 (0·06)
0·95 (0·08)
10·53 (0·82)
0·25 (0·02)
2·55 (0·33)

37·23 (1·04)

numbers by 2·5 times in 2003 (t = 1·97, P = 0·07) and

June 2004

2·0 times in 2004 (t = 3·22, P < 0·01) in the first experi-

ment (Fig. 3a). In the second experiment, the seedling
enhancement ratio as a result of adding seeds was
38·24 (1·04)

3·8 in May, 5·2 in June, 5·5 in July, 6·6 in August and

September 2003 May 2004

17·9 in October (t = 6·94, 6·48, 3·60, 4·43 and 2·14,

respectively; P-values < 0·01 in May–July, P = 0·05 in
October; Fig. 3a). Overall, seedling numbers decreased
Proportion exotic biomass (0·4 m−2)

Grazed (n = 16)

significantly between June and October (Table 4 and

Percentage water 38·92 (1·04)

Fig. 3a).
Litter and standing dead (m−2)

Proportion C4 grass (0·4 m−2)

Proportion exotic S (0·4 m−2)

Grazing alone, without seed additions, did not increase

Proportion grass (0·4 m−2)

number of seedlings 0·1 m−2 for either seed addition

© 2006 The Authors. experiment (Fig. 3a) but grazing conditionally affected
Alpha 1/D (0·4 m−2)
Alpha E (0·4 m−2)
Alpha S (0·4 m−2)

Journal compilation
seedling enhancement. The seedling enhancement
© 2006 British
Biomass (m−2)

Ecological Society,
effect did not differ between grazed and exclosed plots
Journal of Applied in the first experiment but was on average 1·4 times as
Ecology large in grazed than exclosed plots in the second
(Table 4 and Fig. 3b). We did not find a significant
 7

Ecology
© 2006 British
in restorations

Journal of Applied
Native ungulates

Ecological Society,
and seed additions

Journal compilation
© 2006 The Authors.
Table 3. Mean numbers of seedlings m−2 counted in the first and second seed addition experiments. Species listed comprised approximately 90% of those counted. Added species are denoted with an asterisk (*)
and exotic species are in bold (Eilers & Roosa 1994)

2003 2004

Grazed, no seed Grazed, seed Exclosed, no seed Exclosed, seed Grazed, no seed Grazed, seed Exclosed, no seed Exclosed, seed
Scientific name Common name (n = 16) (n = 16) (n = 8) (n = 8) (n = 16) (n = 16) (n = 8) (n = 8)

Seed addition 1
Ambrosia artemesiifolia Common ragweed 24·1 (20·1) 11·6 (5·2) 6·9 (4·5) 3·1 (2·5) 2·0 (0·9) 1·3 (0·9) 0·5 (0·5) 0·5 (0·5)
Aster pilosus Hairy aster 5·3 (2·9) 6·3 (3·7) 13·1 (11·1) 5·0 (3·4) 9·4 (5·5) 9·5 (7·3) 13·0 (9·0) 3·0 (1·7)
Chamaecrista fasciculata* Partridge pea 4·4 (1·6) 5·3 (1·7) 3·8 (2·6) 2·5 (1·3) 6·4 (2·9) 3·6 (1·3) 3·3 (1·8) 7·3 (2·4)
Conyza canadensis Horseweed 3·4 (1·8) 3·4 (1·3) 5·0 (2·7) 5·0 (2·5) 6·4 (2·9) 6·9 (4·8) 6·0 (4·3) 0·8 (0·5)
Daucus carota Queen Anne’s lace 25·9 (15·0) 18·8 (6·5) 11·9 (5·5) 66·9 (46·2) 17·6 (10·1) 22·4 (9·7) 14·3 (6·7) 47·0 (27·1)
Lespedeza capitata* Round-headed bush clover 0·0 (0·0) 6·6 (2·5) 0·0 (0·0) 15·0 (6·9) 0·0 (0·0) 4·9 (1·8) 1·5 (1·0) 4·8 (2·1)
Monarda fistulosa* Wild bergamot 2·5 (1·8) 6·6 (2·2) 5·0 (3·3) 10·6 (7·2) 2·1 (1·1) 15·5 (4·7) 7·8 (5·4) 8·5 (3·2)
Taraxacum officinale Dandelion 30·6 (11·1) 30·6 (7·3) 30·0 (7·8) 33·8 (11·3) 87·3 (26·5) 80·1 (19·2) 59·8 (18·4) 54·8 (17·0)
Other 15·3 (7·3) 15·0 (35·0) 9·4 (3·9) 14·4 (5·7) 8·3 (2·6) 5·6 (1·8) 9·3 (4·2) 8·5 (3·6)
Total number of seedlings 111·6 (35·6) 104·1 (14·8) 85·0 (15·9) 156·3 (50·7) 139·4 (34·5) 149·8 (35·7) 115·3 (28·8) 135·0 (39·3)
Seed addition 2
Aster pilosus Hairy aster 9·4 (5·5) 6·9 (3·0) 14·0 (8·8) 13·3 (5·2)
Chamaecrista fasciculata* Partridge pea 6·4 (2·9) 34·9 (7·6) 3·3 (1·8) 22·0 (4·9)
Conyza canadensis Horseweed 6·4 (5·1) 3·6 (1·8) 6·0 (4·3) 2·8 (0·9)
Dalea purpurea* Purple prairie clover 0·0 (0·0) 18·4 (4·2) 0·0 (0·0) 12·5 (4·0)
Daucus carota Queen Anne’s lace 17·6 (10·1) 39·9 (14·6) 14·3 (6·7) 25·8 (8·6)
Lespedeza capitata* Round-headed bush clover 0·0 (0·0) 96·9 (9·6) 1·5 (1·0) 64·5 (16·8)
Monarda fistulosa* Wild bergamot 2·1 (1·1) 9·6 (2·5) 7·8 (5·4) 7·3 (3·1)
Taraxacum officinale Dandelion 87·3 (26·5) 100·6 (26·5) 59·8 (18·4) 70·0 (35·9)
Other 10·1 (2·6) 59·5 (9·7) 8·8 (3·3) 40·5 (10·3)
Total numbers of seedlings 139·3 (34·5) 351·9 (50·5) 115·3 (28·8) 246·0 (67·3)
8

Table 4.  results (F-values) for seedling enhancement (increase in seedling numbers with seed additions) and seedling numbers between grazed plots and plots exclosed from grazing in a tallgrass prairie

Exotics
L. M. Martin &

0·4
2·1
0·2
B. J. Wilsey

Second experiment seedling numbers

Evenness

6·5*
1·0
1·1
Richness

54·0**
1·8
1·0
Diversity

60·2**
0·3
0·0
Exotics

7·8**
2·0
0·2

0·2
0·4
0·1
Evenness

1·0
0·2
0·9
0·7
0·0
0·4
Richness

9·3**
First experiment seedling numbers

5·1*

1·5
0·2

0·0
0·2
Diversity

10·7**
4·0†

1·2
0·7

0·1
0·3
Num/Den d.f.
restoration. Significance is indicated by †P between 0·06 and 0·1, *P between 0·02 and 0·05 and **P ≤ 0·01

1/22
1/22

1/44

2/44
1/44
1/15
Fig. 3. The (a) number of seedlings 0·1 m−2 in grazed and
exclosed plots when seeds were added or were not and (b)
effects of grazing on seedling enhancement for two experi- Seedling enhancement
mental seed addition trials in a tallgrass prairie restoration
(n = 16 for grazed; n = 8 for ungrazed). The first seed addition
Second experiment

experiment is presented before the break in the x axis and the

second experiment is after the break. Vertical bars are ± 1 SE.

7·3**
6·3*

0·5
relationship with GI in either experiment (first experi-
d.f.

4
15

ment, linear effects F1,13 = 0·01, P = 0·93, quadratic 88

effects F1,13 = 1·68, P = 0·22; second experiment, linear
Seedling enhancement

effects F1,13 = 0·63, P = 0·44, quadratic effects F1,13 =

2·14, P = 0·17).
Path analysis from both experiments indicated that
First experiment

biomass m−2 was negatively related to light and soil

water availability, and that water was positively related
0·02

0·5
0·6

to seedling enhancement more regularly and strongly

than light (Fig. 4 and Table 5). The indirect pathway
d.f.

15
1
1

22

indicated that GI was positively related to NPP in the

first (significant for 2004 only) and second experiments,
Time × grazed × seed addition

but NPP never significantly explained seedling enhance-

ment beyond effects of light and water (Fig. 4 and Table 5).
Grazed × seed addition

© 2006 The Authors.

Journal compilation
© 2006 British Exotic species seedling emergence
Time × grazed
Seed addition

Error (time)

Ecological Society,
Journal of Applied Exotic and cosmopolitan weed seedling emergence
Grazed
Source

were not clearly affected by treatments in either experi-

Error
Time

Ecology
ment (Table 4). Exotics such as Taraxacum officinale
9 mean number of exotics was not related to mean GI
Native ungulates for either experiment (first, linear, exotics F1,13 = 0·07,
and seed additions P = 0·79; quadratic, exotics F 1,13 = 2·08, P = 0·17;
in restorations second, linear, F1,13 = 0·59, P = 0·45, quadratic,
F1,13 = 0·23, P = 0·64).

  

Fig. 4. Path analysis diagram that tests direct effects of
grazing intensity on seedling enhancement through biomass,
light and water, and indirect effects of grazing intensity on
seedling enhancement through NPP effects in a tallgrass
prairie restoration. Ten and 25 rare prairie species were added
in plots inside and outside grazing exclosures in two separate
experiments.
and Daucus carota were among the most abundant
species in both experiments, and cosmopolitan weeds
Aster pilosus and Ambrosia artemisiifolia were also
abundant in the first experiment (Table 3). The mean
number of exotic seedlings did not significantly differ
between grazed and exclosed plots in either experiment
[mean (SE) number of exotics 0·1 m−2, n = 48; first
experiment 2003, grazed 1·51 (0·28), exclosed, 1·72
(0·31); 2004, grazed 2·03 (0·28), exclosed 1·91 (0·31);
second experiment, grazed 2·19 (0·29), exclosed 1·92
(0·31); Table 4]. Grazing effects did not interact with
seed additions in either experiment (Table 4). The
Seed additions increased seedling diversity and richness
in both experiments and slightly decreased seedling
evenness in the second experiment. Mean diversity in
the first experiment was 1·2 times as great with seed
additions in 2003 and 2004, but this difference was only
marginally significant [mean (SE) 1/D 0·1 m−2, n = 48;
2003, 2·76 (0·22), control 2·24 (0·22); 2004, 2·10 (0·22),
control 1·82 (0·22); Table 4]. Mean diversity in the
second experiment was 1·9 times as great in the seed
addition subplots than in controls [mean (SE) 1/D
0·1 m−2, n = 48; addition 3·37 (0·20), control 1·82 (0·20);
Table 4]. Mean richness in the first experiment was 1·3
and 1·2 times as great in seed addition than control sub-
plots in 2003 and 2004, respectively, and was 2·3 times
as great in addition subplots in the second experiment
[mean (SE) S 0·1 m−2, n = 48; first experiment, 2003,
addition 3·81 (0·32), control 2·88 (0·32); 2004, addition
2·96 (0·32), control 2·53 (0·32); second experiment,
addition 5·82 (0·37), control 2·52 (0·37); Table 4]. Mean
evenness did not differ in the first experiment but was
slightly lower in seed addition subplots in the second
[mean (SE) E 0·1 m−2, n = 48; first 2003, addition 0·75
(0·03), control 0·77 (0·03); 2004, addition 0·74 (0·03),
control 0·72 (0·03); second, addition 0·65 (0·03), con-
trol 0·72 (0·03); Table 4].
Seedling species diversity did not differ between
grazed and exclosed plots for either experiment [mean
(SE) for grazed and exclosed, respectively, n = 48; first
experiment, diversity 2003 2·47 (0·19), 2·53 (0·25), 2004

Table 5. Path analysis results to determine direct effects of grazing intensity on biomass, biomass on light and water, and light and
water on seedling enhancement, and indirect effects of grazing intensity on NPP and NPP on seedling enhancement for the first
and second seed addition experiment. Ten (first) or 25 (second) rare prairie species were added to plots. Numbers represent
standardized regression coefficients. Significance is indicated by *P between 0·02 and 0·05 and **P ≤ 0·01

First experiment Second experiment

Variable Second half 2003 First half 2004 Second half 2004 First half 2004 Second half 2004

Direct effects
GI → biomass 0·00 0·05 − 0·27 0·05 − 0·27
Biomass → light − 0·73** 0·04 − 0·65** 0·04 − 0·65**
Biomass → water − 0·61** − 0·24 − 0·65** − 0·24 − 0·65**
Light → enhancement 0·30 0·15 − 0·16 0·16 0·34*
Water → enhancement − 0·23 0·51** 0·43* 0·56** 0·53**
Indirect effects
GI → NPP 0·09 0·75** 0·55* 0·75** 0·55**
© 2006 The Authors. NPP → enhancement 0·11 − 0·19 − 0·16 − 0·06 − 0·08
Journal compilation Unobserved
o1 → biomass 1·00** 1·00** 0·96** 1·00** 0·96**
© 2006 British
o2 → light 0·68** 1·00** 0·76** 1·00** 0·76**
Ecological Society,
o3 → enhancement 0·95** 0·82** 0·91** 0·81** 0·68**
Journal of Applied
o4 → water 0·79** 0·97** 0·76** 0·97** 0·76**
Ecology
o5 → NPP 1·00** 0·66** 0·83** 0·66** 0·83**
10 1·88 (0·19), 2·04 (0·25), richness 2003 3·28 (0·29), 3·41
L. M. Martin & (0·36), 2004 2·68 (0·29), 2·81 (0·36), evenness 2003 0·75
B. J. Wilsey (0·03), 0·77 (0·04), 2004 0·71 (0·03), 0·75 (0·04); second
experiment diversity 2·58 (0·19), 2·61 (0·22), richness
4·39 (0·33), 3·95 (0·42), evenness 0·66 (0·02), 0·71
(0·04); all at the 0·1 m2 scale; Table 4]. Diversity
enhancement declined with GI in the first experiment
(linear effect F1,13 = 5·05, P = 0·04; quadratic effect
F1,13 = 2·29, P = 0·15; data not shown) and was quad-
ratically related to GI in the second experiment (F1,13 =
6·02, P = 0·03; quadratic equation y = 2·4 ± 11·3x +
15·5x2) with an outlier included, and unrelated with
an outlier excluded (linear effect F1,13 = 3·22, P = 0·10,
r = 0·45, slope = 1·83; quadratic effect F1,13 = 0·53,
P = 0·48; data not shown). The evenness enhancement
effect was negatively related to GI in the first experi-
ment (F1,13 = 4·8, P = 0·05, r = 0·53, slope = −0·53) only.
Discussion
Previously, we found that conventional prairie restora-
tion at the study site was able to restore common native
species but not species diversity of nearby prairie
remnants (Martin, Moloney & Wilsey 2005). Here we
tested hypotheses regarding why diversity was lower.
Our results suggest that seedling emergence in low
diversity restorations is seed limited but that native
ungulates can sometimes increase emergence as well.
Seedling enhancement increased with water and light
availability, which suggests that, when grazing is enhanc-
ing emergence, the mechanism may be associated with
grazers having a direct effect on water and light. A
combination of seed additions and grazing led to the
highest amount of seedling emergence, but this result
was conditional, i.e. it was found only in the second
trial and year. However, grazing alone did not increase
seedling emergence in either trial.
Biomass and NPP, general indicators of resource
uptake in grasslands, are usually affected by grazing
(Semmartin & Oesterheld 1996). We found that total
biomass declined with grazing intensity, and water and
light availability were higher when biomass was lower,
which suggests that grazing decreased biomass enough
to increase resource availability. However, ANPP was
also higher with grazing, suggesting that defoliated plants
were readily recovering from defoliation and utilizing
available resources (Knapp et al. 1999; Wilsey et al.
2002). Nevertheless, increased levels of ANPP did
not appear to have effects above and beyond those
correlated with water and light availability in reducing
seedling enhancement.
© 2006 The Authors.  
Journal compilation
Seedling emergence was limited by seed availability in
© 2006 British
Ecological Society,
these tallgrass prairie restorations, which suggests that
Journal of Applied restorations are similar to many old fields in their lack
Ecology of propagule availability (Tilman 1997; Zobel et al.
2000; Pywell et al. 2002). Many tallgrass prairie restora-
tions are initiated by harvesting seeds from remnants
in the autumn (Polley, Derner & Wilsey 2005), when C4
grass seed is most abundant relative to other species.
Evidence from this study suggests that seedling
emergence of rare forbs is very low nearly 10 years after
initial seeding, and that adding seeds of rare forb and
grass species that are typically lacking in restorations
can increase seedling emergence, the first step in
recruiting species into the community.
Our finding that seed additions and grazing com-
bined could increase seedling emergence suggests that
grazing mammals might increase seedling recruitment
in some situations. Turnbull, Crawley & Rees (2000)
found that, overall, a combination of adding seeds and
inducing disturbance to reduce dominant vegetation
was most important for recruitment. Rhinanthus minor,
a parasitic plant, also increased seedling recruitment
of added species by reducing competitive effects of
dominant vegetation (Pywell et al. 2004). However, the
effect of grazing on seedling emergence with seed addi-
tions in the restoration was conditional. This condi-
tionality may have been the result of very different
weather between years, but our design could not deter-
mine this definitively. Although grazing combined with
seed additions conditionally improved seedling emer-
gence, we did not find that emergence was quadratically
related to grazing intensity, as predicted by the inter-
mediate disturbance hypothesis. Knapp et al. (1999)
proposed that target grazing intensity in intact tallgrass
prairies should be about 25% of annual above-ground
primary production, based on historic grazing intensi-
ties. We observed grazing intensities in the restoration
that were sometimes double that estimate. In our study,
it appears that grazing had an increasingly beneficial
(i.e. linear) effect on seedling emergence when seeds
were added.
Increases in exotic or cosmopolitan weed species may
negatively impact diversity and are a primary concern
in grazed grasslands (Smith & Knapp 1999; Hulme &
Bremner 2006). Grazing, which is utilized in manage-
ment of both intact grasslands and restorations
(Collins et al. 1998; Knapp et al. 1999), could increase
weeds by the same mechanisms that increase native
plant recruitment (Smith & Knapp 1999). In contrast,
high dominance may enhance seedling emergence in
some grasslands (Wilsey & Polley 2002; Smith et al.
2004). We found no relationships with grazing on
exotics, suggesting that grazing may not be important
to exotic recruitment in these restored grassland
communities.
   

Diversity components responded differently to seed
additions and grazing. With grazing, we found no
changes in species richness in the vegetation after
1.5 years of grazing, but evenness increased linearly
with grazing intensity. An increase in evenness can be
11 associated with a decrease in dominance from ungulate
Native ungulates activities (Hartnett, Hickman & Walter 1996). How-
and seed additions ever, grazers may not be able to recruit new species and
in restorations enhance richness if rare plant species are not available
either in the seed bank or as vegetative propagules,
which appears to be the case in this restoration. Seed
additions, on the other hand, tended to increase rich-
ness but had a smaller negative effect on evenness.
It is important to point out that we only tested for
seed limitation with the emerging seedling community.
Although seedling emergence is a crucial step towards
recruitment into a community, seedlings must establish
viable populations before they will influence diversity
in the longer term. Wilsey & Polley (2003) found that,
even when seedling emergence was high, plant diversity
was unchanged because of low seedling survival in a
Texas grassland. It is too early to estimate establish-
ment success, and longer term monitoring is needed to
test the hypothesis that seed additions and grazing will
increase diversity of vegetation in the long term.
  

Seedling numbers of rare species increased greatly when
seeds were added, suggesting that these restorations
were severely seed limited. Grazers increased water and
light availability by decreasing above-ground biomass,
and water increased seedling enhancement more regu-
larly and strongly than did light. Grazing increased
ANPP, but ANPP did not explain seedling emergence
above and beyond water and light effects. Under
certain conditions, seedling emergence was positively
influenced by grazing when seeds were added. However,
grazing by itself did not increase seedling emergence,
probably because seeds of rare species were not
available to emerge. These results suggest that it may be
advantageous to mimic positive effects of native un-
gulates in restorations when seeds or propagules of rare
species are available to emerge. If propagule availability
is low, positive effects of grazing alone will not increase
seedling emergence. Improving seedling emergence in
restorations may therefore require adding seeds even
after dominant plants are established, while simultane-
ously mimicking grazing effects to increase light and
water availability. In contrast to management strategies
in intact grasslands (Collins et al. 1998), utilizing
grazing without propagules may not improve seedling
emergence in restorations. However, longer term moni-
toring is necessary to determine if seed additions and
grazing promote long-term plant diversity.
© 2006 The Authors. Acknowledgements
Journal compilation
We thank Pauline Drobney at Neal Smith NWR,
© 2006 British
Ecological Society,
Andrea Blong, Brennan Dolan, Dan Haug, and David
Journal of Applied Losure for help in the field, and Tim Dickson and
Ecology two anonymous referees for comments on an earlier
version of this manuscript.
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Received 31 October 2005; final copy received 28 May 2006
Editor: Phil Hulme