Download Regional adaptation improves the performance of grassland plant

Basic and Applied Ecology 13 (2012) 551–559
Regional adaptation improves the performance of grassland plant
communities
Karoline Weißhuhna,∗ , Daniel Pratib , Markus Fischerb , Harald Augea
a
UFZ – Helmholtz Centre for Environmental Research, Department of Community Ecology, Theodor-Lieser-Straße 4, D-06120 Halle, Germany
Institute of Plant Sciences and Oeschger Centre for Climate Change Research, University of Bern, Altenbergrain 21, CH-3013 Bern,
Switzerland
b
Received 22 January 2012; accepted 15 July 2012
Abstract
Many plant species are adapted locally or regionally. Whether such individual species performance translates into effects at
community and ecosystem levels has rarely been tested. Such tests are crucial, however, to predict ecosystem consequences
of sowing seed mixtures for grassland restoration or hay production. We compared the performance of replicated sown plant
communities of regional origin with the performance of four foreign communities consisting of the same grassland species but
originating from distances up to 890 km from our experimental site. The regional communities performed better than foreign
communities in plant cover and diversity but not in aboveground biomass production. Additionally, in communities based
on regional seeds fewer unsown species occurred and less bare ground was left open for erosion. Variation in community
performance among source regions was related to climatic differences rather than to geographic distance to source regions.
Individual species performance only partly explained community patterns, highlighting the importance of community level
experiments. Our results suggest that the use of regional seeds represents an important approach to improve sown managed
grasslands.
Zusammenfassung
Viele Pflanzenarten sind lokal oder regional angepasst. Ob sich diese Entwicklung einzelner Arten in Effekte auf der Ebene von
Gemeinschaften oder Ökosystemen umsetzt, ist kaum untersucht. Dies wäre aber wichtig, um die Konsequenzen der Verwendung
von Saatgutmischungen für Heuwiesen oder die Restaurierung von Grasland vorherzusagen. Wir verglichen angesäte replizierte
Graslandgemeinschaften regionaler Herkunft mit vier Gemeinschaften gleicher Artenzusammensetzung, aber fremder Herkunft
aus Entfernungen bis zu 890 km von der Experimentalfläche. Die Gemeinschaften regionaler Herkunft entwickelten eine höhere
Pflanzendeckung und Diversität als die fremden Herkünfte, produzierten aber nicht mehr oberirdische Biomasse. Außerdem
traten in den Gemeinschaften regionaler Herkunft weniger nicht eingesäte Pflanzenarten und für Erosion besonders anfälliger
unbedeckter Boden auf. Unterschiede zwischen Gemeinschaften unterschiedlicher Herkunft hingen eher mit dem Herkunftsklima zusammen als mit der geografischen Distanz zur Experimentalfläche. Das Verhalten der einzelnen Arten konnte die
Gemeinschaftsmuster nur teilweise erklären, was die Wichtigkeit von Experimenten auf der Gemeinschaftsebene unterstreicht.
Unsere Ergebnisse legen nahe, dass der Einsatz von regionalem Saatgut einen wichtigen Ansatz für die Verbesserung von
angesätem Wirtschaftsgrünland darstellt.
© 2012 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
Keywords: Central Europe; Climate; Community ecology; Grassland management; Multi-species transplant experiment
∗ Corresponding
author. Tel.: +49 345 558 5302; fax: +49 345 558 5329.
E-mail address: [email protected] (K. Weißhuhn).
1439-1791/$ – see front matter © 2012 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
http://dx.doi.org/10.1016/j.baae.2012.07.004
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K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
Introduction
Materials and methods
Many plant species are locally or regionally adapted to
their environment as demonstrated by reciprocal transplant
experiments (Leimu & Fischer 2008). An advantage of local
or regional plants and seeds mainly results from adaptation
to factors such as climate, soil or land use (Macel et al. 2007;
Raabova, Muenzbergova, & Fischer 2007). If plant species
within one provenance are adapted to each other, this may also
modify their interactions (Bischoff et al. 2006), which in turn
may also affect ecosystem properties. While transplant studies showed the importance of local or regional provenances
for the performance of individual species, much less is known
about whether such adaptation also translates into community
effects. If this is the case, the performance of communities
of identical species composition may differ between communities composed of local provenances and communities
composed of provenances from further away. Additionally,
such within-species variation may well influence ecosystem
processes (Whitlock, Grime, & Burke 2010). To our knowledge, however, experiments addressing the community or
ecosystem level effect of geographic variation in species’
origin are lacking.
In restoration ecology the use of regionally adapted seed
material has been discussed controversially (Hamilton 2001;
Wilkinson 2001). Whereas local seed material has been
debated as the best choice in forestry, a similar acceptance
for grassland is largely lacking, as evidenced for instance
by the absence of seed exchange regulations by the European Union (see Vander Mijnsbrugge, Bischoff, & Smith
2010). This underlines the need for experiments investigating whether regional adaptation of plant populations
also translates into effects at the level of communities and
ecosystems.
Regional adaptation may affect plant performance at the
community level at an early stage of establishment when
germination rates differ between seed origins (Keller &
Kollmann 1999). At later stages plant performance may be
more related to differences in climatic conditions rather than
geographic distance (Leimu & Fischer 2008), although distance could be a good predictor for adaptive divergence
of populations (Joshi et al. 2001; Becker, Colling, Dostal,
Jakobsson, & Matthies 2006). If community performance and
properties would show patterns of regional adaptation, further
evidence for the need to use local and regional seed material
would be provided.
We use a field experiment with commercially available
grassland seed mixtures to test (i) whether replicated plant
communities of regional provenance establish better than
communities composed of foreign provenances of the same
species, (ii) whether this advantage of the regional community translates into improved ecosystem properties such
as productivity, weed suppression and soil protection and
(iii) whether variation in performance among provenances
is related to geographic distance or climatic conditions in the
source regions of the seed material.
Study system
We used seed mixtures of 11 grassland plant species from
different provenances. These species are commonly used in
commercial seed mixtures for grassland restoration in mesic
habitats and include two grasses, eight non-legume forbs and
one legume (for species see Table 1, first column).
We selected five European regions and purchased seeds
of each of the 11 species from commercial suppliers in each
region (see Appendix A: Table S1 Central Germany (CG),
Baden-Württemberg (BW) and Bavarian prealps (BA) in Germany, north-eastern Switzerland (CH) and southern Great
Britain (GB). The suppliers guaranteed regional origin of the
seed material and that species had not been changed by breeding because commercial production of plants can impose
selection on some plant traits. Although the sampling site
CG is located 90 km from our field experimental site, we consider this provenance as regional because the two sites belong
to the same floristic region “Thüringisch-Fränkisches Mittelgebirge” (Meynen & Schmithüsen 1953; Rüdiger Prasse,
University of Hannover, personal communication). In the following, we will apply the term “regional adaptation” (Ridley
& Ellstrand 2010) to distinguish adaptive divergence of plant
performance among different regions from local adaptation
occuring at small spatial scales, i.e. within regions or habitats.
Since seeds of Trifolium pratense were not available for
the provenance BA, we replaced it with the BW provenance
of this species, which is geographically close to BA. For the
GB provenance, Hypochaeris radicata had been delivered
instead of Leontodon autumnalis L., which we noticed only
at the first vegetation survey. However, as H. radicata and L.
autumnalis are closely related to each other, and of the same
growth form, we present results including British H. radicata.
For each provenance, we mixed seeds of all species according to recommendations for grassland establishment so that
legume and non-legume herbs together contributed 50% to
total seed mass, while grasses contributed the remaining 50%.
Each species was represented by approximately the same
numbers of seeds in each group.
Field experiment
The experiment was set up on an abandoned arable field
with carbonate-free and nutrient-poor soil characterised by
pH 5.4 (H2 O), C/N ratio of 10.45 and plant-available P of
98 mg/kg (double lactate extraction). The field was ploughed
in autumn 2007 and harrowed in spring 2008. Thereafter,
we established 60 plots of 2.5 m × 2.5 m with an inter-plot
distance of 1.5 m in a randomised block design with ten
blocks, each containing all six treatments. In each block the
treatments involved sowing seeds from one of the five provenances to a density of 4 g m−2 , and an unseeded plot to assess
spontaneously germinating species. In April 2008, the seeds
were mixed with soy granulate (20 g m−2 ) to allow an even
Species
2008
2009
Block (DF = 9, 36)
Grasses
Anthoxanthum odoratum L.
Holcus lanatus L.
Non-legume herbs
Achillea millefolium L.
Campanula rotundifolia L.
Centaurea jacea L.
Knautia arvensis (L.) J.M. COULT
Leontodon autumnalis L.,
Hypochaeris radicata L.
Leucanthemum vulgaris LAM
Plantago lanceolata L.
Silene vulgaris (MOENCH) GARCKE
Legume herb
Trifolium pratense L.
Provenance (DF = 4, 36)
Block (DF = 9, 36)
Provenance (DF = 4, 36)
SS
SS
SS
F
SS
F
0.056
335.1
0.65
1.69
0.216
1135
5.69***
12.9***
59.85
2.84*
40.02
18.21
0.55
313.1
0.58
348.1
9.833
6.123
80.34
3.308
1.928
0.84
4.29***
1.34
2.21
28.45
55.84
6.566
1.365
F
F
0.84
0.69
70.64
201.3
9.56*** ↑
11.3***
981.4
2.18*
553.1
2.77***
21.1*** ↑
100.5
0.65
853.6
12.5*** ↑
46.1*** ↑
108.0
0.86
3924
70.4*** ↑
8.78***
6.72*** ↓
5.97***
156.8
110.9
0.570
0.66
1.75
0.48
3840
269.2
1.942
36.5***
9.57***
3.71***
3.53***
163.1
1.97
56.96
1.54***
4.27*** ↑
14.12
27.86
K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
Table 1. Summary of ANOVA of cover by different sown plant species in our experiment with sown grassland plant communities of different seed origins. Significance levels are corrected
for multiple comparisons. For Campanula rotundifolia and Knautia arvensis we did not have enough data points for this analysis. Where the contrast regional vs. foreign provenances was
significant, ↑ indicates better and ↓ worse performance of the regional provenance (*p < 0.05; ***p < 0.001).
553
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K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
distribution of plant seeds, spread by hand and gently rolled
onto the soil. The unseeded plots were treated in the same
way except that soy granulate was spread without seeds. Seed
material of all species in all provenances was germinable as
tested in the lab. Plots were not weeded because the interaction of sown with unsown species was considered to be an
integral part of the experiment. In the unseeded plots none of
the sown species occurred, hence we conclude that the target
plant species indeed emerged from sown seeds in sown plots.
In July 2008 and June 2009 we carried out vegetation
surveys on a 1 m2 area within each plot by estimating the
cover of all vascular plant species visually using a modified
Londo scale (Londo, 1984). To measure above-ground productivity we clipped all vegetation 2 cm above ground on a
50 cm × 50 cm within each plot in August 2008 and July 2009
when vegetation surveys were done. Afterwards the whole
field was mown according to the common management practice of the surrounding grasslands and plant material was
removed. The clipped biomass was dried at 60 ◦ C for 72 h
and weighed. As a proxy for exposure to soil erosion, we estimated the proportion of bare ground according to the Londo
scale. The monthly climate variables precipitation and temperature were close to the long term means (see Appendix A:
Table S1). Although precipitation in May 2008 after sowing
was low, the high precipitation January to March resulted in
sustainedly high soil moisture.
Climatic conditions differed substantially among the
regions of the five provenances (see Appendix A: Table S1).
To characterise them, we calculated mean annual temperature
to depict the North to South gradient, annual temperature
range to describe the continentality gradient, and accumulated growing degree days (the sum of mean day temperatures
of days warmer than 5 ◦ C) to describe the growing season
length. Because the measurements started in June, data from
January to June from 1971 to 2000 (Mitchell, Carter, Jones,
Hulme, & New 2004; New, Hulme, & Jones 1999) were used
for our calculations of climate variables, representing interpolations of a 10 grid averaged over all grid cells from which
the seed material had been sampled. This long term data may
represent the climatic conditions plants had adapted to in their
home regions. We calculated Pearson correlation coefficients
between these climatic variables and the mean plant cover of
each seed provenance for both years and corrected for multiple comparisons. All analyses were done with the statistical
software JMP8 (SAS Institute Inc., Cary, NC, USA).
Statistical analysis
Shannon diversity H = −pi × log(pi ) was based on cover
data of sown species, where pi is the frequency of species i.
Due to the same number of sown species and the small number of extinctions, differences in species richness were very
small. Therefore, Shannon evenness was similar to Shannon diversity and is not presented. As we were interested
in the performance of the sown communities, we calculated
diversities for the sown species only. We performed separate ANOVAs for cover of sown, unsown and all species
(cover sown + cover unsown species), bare ground, biomass
and diversity with block and provenance as factors using
type I sums of squares. We treated provenance as a fixed
effect because provenances were chosen according to climatic conditions and geographic distance to our field site. We
defined the contrast regional vs. foreign to test whether the
regional provenance (CG) performed better than the foreign
provenances. Within each year, levels of significance were
corrected for multiple comparisons (Benjamini–Hochberg
procedure; Verhoeven, Simonsen, & McIntyre 2005). Just
the terms, which were significant after correction, were
considered as significant in the discussion. To fulfil the preconditions for ANOVA, data were Box–Cox-transformed
prior to analyses. The same model was also calculated for
each species and year separately except for Campanula
rotundifolia and Knautia arvensis, which occurred very
rarely in the experimental communities.
Fig. 1. Plant cover of sown and unsown species in 2008 and 2009
for replicated regional (CG) and increasingly distant foreign plant
communities (from BW to GB) sown at one experimental site.
Stacked bars represent means of sown species, black circles represent means (±standard error) of unsown species. Campanula
rotundifolia occurred rarely and is not shown. CG = Central Germany, BW = Northern Baden-Württemberg, BA = Bavarian prealps,
CH = North-eastern Switzerland, GB = Southern Great Britain.
K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
555
Table 2. Summaries of ANOVA of annual measures of community performance in our experiment with sown grassland plant communities
of different seed origin. Where the contrast regional vs. foreign provenance performance was significant, ↑ indicates higher, and ↓ lower
performance of the regional than for the foreign communities (*p < 0.05; **p < 0.01; ***p < 0.001).
Source
Cover sown species [%]
Block
Provenance
Regional vs. foreign
Residuals
Residuals
R2
Cover unsown species [%]
Block
Provenance
Regional vs. foreign
Residual provenance
Residuals
R2
Cover all species [%]
Block
Provenance
Regional vs. foreign
Residuals
Residuals
R2
Shannon diversity index
Block
Provenance
Regional vs. foreign
Residual provenance
Residuals
R2
Bare ground [%]
Block
Provenance
Regional vs. foreign
Residual provenance
Residuals
R2
Aboveground biomass [kg/m2 ]
Block
Provenance
Regional vs. foreign
Residual provenance
Residuals
R2
2008
2009
DF
SS
F
DF
SS
F
9
4
1
3
36
0.64
0.17
0.21
0.07
0.14
0.21
3.13*
8.75***
11.6** ↑
9
4
1
3
36
0.54
0.73
0.43
0.31
0.12
0.96
3.02*
4.00*
11.7* ↑
9
4
1
3
36
0.59
0.41
0.18
0.07
0.11
0.41
3.98**
4.03*
6.35* ↓
9
4
1
3
36
0.49
0.00
0.06
0.05
0.01
0.06
0.30*
7.88***
27.1*** ↓
9
4
1
3
36
0.59
0.52
0.03
0.00
0.03
0.39
5.37**
0.75
0.06
9
4
1
3
36
0.49
0.86
0.20
0.14
0.06
1.12
3.07*
1.62
4.43
9
4
1
3
36
0.36
0.05
0.09
0.00
0.09
0.24
0.78
3.38*
0.17
9
4
1
3
36
0.58
0.02
0.24
0.04
0.20
0.18
0.41
11.8***
7.09* ↑
9
4
1
3
36
0.43
0.04
0.11
0.01
0.10
0.20
0.74
5.10**
2.23
9
4
1
3
36
0.47
0.20
0.13
0.09
0.04
0.38
2.11
3.11*
8.69* ↓
9
4
1
3
36
0.46
0.10
0.03
0.01
0.02
0.16
2.52*
1.88
3.15
9
4
1
3
36
0.40
0.10
0.03
0.01
0.02
0.21
1.96
1.52
1.53
Results
Provenance effects on community performance
and properties
The five provenances varied in their establishment success
and performance (Fig. 1, Table 2). Establishment in 2008
was significantly higher for the regional “home” provenance
with a plant cover of sown species of 38 ± 3.3% compared
with 28 ± 1.6% averaged over the foreign provenances. Total
cover of all species (sown plus unsown) did not differ between
communities of different sown provenances.
By the second year the Shannon diversity H of communities of regional provenance was higher than the
average diversity of the foreign provenances (Fig. 2A,
Table 2). Thus, with increasing time the use of regional seed
material led to a more even distribution of species in the
community.
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K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
Fig. 2. Estimated proportion of the Shannon diversity index H (A)
and bare ground (B) in June for replicated regional plant communities (CG) and increasingly distant foreign comunities (from
BW to GB) sown at one experimental site (means ± standard
error). See Table 2 for contrast analysis of regional vs.
foreign communities. CG = Central Germany, BW = Northern
Baden-Württemberg, BA = Bavarian prealps, CH = North-eastern
Switzerland, GB = Southern Great Britain.
Above-ground community biomass production did not differ among provenances. It should be noted, however, that we
did not separate biomass of sown species and unsown species,
hence these results refer to the total biomass.
In the regional communities fewer unsown species
grew than in foreign communities in both years (Fig. 1)
(cover percentages of unsown species: 2008: regional
28 ± 4.9%, foreign 37 ± 2.6%; 2009: regional 4 ± 0.6%, foreign 11 ± 1.0%), suggesting that suppression of unsown by
sown species in the first year determined their development
in the subsequent year and that unsown species did not play
a major role after the first year.
As a measure of potential risk of soil erosion, we estimated
the proportion of bare ground exposed to erosion after mowing the plots (Fig. 2B). In both years, the proportion of bare
ground was lower in communities of regional provenance
than in communities of foreign provenances (2008: regional
9 ± 2.1%, foreign 14 ± 1.7%; 2009: regional 34 ± 2.2%, foreign 45 ± 2%).
Fig. 3. Relationships between (A) cover by sown plant communities
in 2008 and mean annual temperature in the regions of seed origin,
and (B) cover by sown communities in 2009 and accumulated growing degree days in the regions of seed origin (data 1971–2000, New
et al., 1999; Mitchell et al., 2004). The experimental site had a mean
annual temperature of 6.97 ◦ C and 573 agdd. CG = Central Germany, BW = Northern Baden-Württemberg, BA = Bavarian prealps,
CH = North-eastern Switzerland, GB = Southern Great Britain.
Community performance related to climate and
distance
Variation in plant cover among provenances was best
explained by mean annual temperature in the source region
in 2008 and by the accumulated growing degree days in the
source region in 2009 (Fig. 3, Table 3), but was not related
to geographic distance. Thus, climatic differences among
source regions explained differences in plant performance
better than geographic distance did.
Provenance effects on individual species
performance
Plant establishment in 2008 and performance in 2009 varied among provenances in all nine analysed species (Fig. 1,
Table 1). Contrast analysis comparing regional and foreign provenances within species for each of the two years
revealed higher establisment or performance of the regional
K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
557
Table 3. Correlations between community cover in our experiment with sown grassland-plant communites of different seed origin and climate
variables for the five regions of seed origin (n = 5). Adjusted significance are the significance levels after correction by the Benjamini–Hochberg
procedure (see Methods section; *p < 0.05; **p < 0.01).
Plant cover in 2008
Distance to field site [km]
Mean annual temperature [◦ C]
Range of annual temperature [◦ C]
Accumulated growing degree days January–June
Plant cover in 2009
R
p
Adjusted
significance
R
p
Adjusted
significance
−0.91
−0.97
0.68
−0.63
0.030
0.005
0.205
0.256
ns
*
ns
ns
−0.40
−0.64
−0.02
−0.99
0.507
0.240
0.978
0.0006
ns
ns
ns
**
provenance in six out of 18 cases (9 species × 2 years)
and lower establishment in one case. In the remaining 11
of 18 comparisons, variation in individual-species establishment or performance was idiosyncratic with respect to
seed origin and revealed neither a home-site advantage nor
a disadvantage of the regional provenance. This indicates
that, in contrast to the community as a whole, individualspecies performances did not conform clearly to regional
adaptation.
Discussion
Effects of regional vs. foreign provenances on
community performance
Generally, the better performance of communities of
regional provenance over those of foreign provenance suggests adaptation (Kawecki & Ebert 2004). Although we had
one experimental site only, the explained variance of about
99% of plant cover in the second year by climate variables
in the region of seed origin suggests that plant performance
was mainly determined by climatic differences between plant
origins. Furthermore, it is highly unlikely that data of two
years comparing five provenances would by chance reveal
the consistent result of the best performance of the regional
provenance.
Adaptations of single species to regional factors were
repeatedly shown (e.g. Becker et al. 2006; Raabova et al.
2007), but community response is hard to predict from such
results due to differences among species in presence or
absence of local adaptation (Jones & Hayes 1999; Leimu
& Fischer 2008). In our experiment, regional adaptation of
communities was demonstrated experimentally for the first
time. Since the provenances used originate from one region
but not necessarily from one particular site, the observed
pattern of adaptation is likely to reflect evolutionary adaptation to factors acting at a regional rather than at a local
scale. Such regional effects are known for plant adaptation to abiotic factors such as climate or soil (Macel et al.,
2007). Inter-continental comparisons indicate that adaptation
among species can also occur at such large spatial scales,
e.g. between competitors (Callaway & Aschehoug 2000),
between plants and pathogens, herbivores (Vilà, Maron, &
Marco 2005) or mutualists (Stinson et al. 2006).
If plants of regional provenance perform better on average
than plants of foreign provenances, the chance of individual species failing or dominating at a regional site should
be lower and interactions between plant species may be better balanced. Additionally, a shared co-evolutionary history
may improve resource partitioning and coexistence among
plant species within a community, as suggested by comparing plant communities composed of species native and exotic
to the U.S. (Wilsey, Teaschner, Daneshgar, Isbell, & Polley
2009). Together, this may lead to more efficient resource use
which may provide an explanation for the lower percentage
of bare ground and stronger suppression of unsown species
in communities of regional provenance in our experiment.
Possibly, also different responses to diseases between
plants of regional and foreign origin (Cremieux et al. 2008)
may have contributed to higher performance of regional communities in our experiment. Geographical scales at which
plants could be adapted to each other are difficult to define
and vary among species (Vander Mijnsbrugge et al. 2010).
There is some evidence that a history of coexistence within a
region can affect evolutionary trajectories of species and the
outcome of interspecific interactions (Thorpe, Aschehoug,
Atwater, & Callaway 2011). However, individual species
responses are highly influenced by competitive relationships
and individual stress responses (Grime 1974) which together
shape plant communities. Consequently, community performance appears much more meaningful than studies of single
species or monocultures for addressing determinants of grassland plant community performance.
Provenance effects on ecosystem properties
Comparing communities comprised either of native or of
invasive plants indicated that a common evolutionary history of species may affect ecosystem processes (Wilsey et al.
2009). Our study suggests that similar effects may occur at the
level of regional provenances of species composing communities. To our knowledge, ours is the first experiment showing
regional adaptation of a set of species at the community
level. Several ecosystem properties, including suppression
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K. Weißhuhn et al. / Basic and Applied Ecology 13 (2012) 551–559
of unsown species, prevention of soil erosion, and Shannon
diversity, were higher for communities composed of regional
provenances than for those composed of foreign provenances.
Previous studies which suggested the use of native and locally
adapted plants to improve the ecosystem relied heavily on
results for and effects of individual and particularly influential species (Fiedler, Landis, & Wratten 2008). We support
the recommendation based on our community experiment
and argue that more studies should be conducted with entire
communities, or at least subsets thereof, to assess ecosystem
consequences of local and regional seed material.
Central European grasslands are characterised by a high
frequency of ploughing and re-seeding and a high proportion
is managed by mowing. This bears the risks of exposing large
proportions of bare soil to potential erosion. In our experiment, less open ground was exposed to erosion in regional
communities than in foreign ones. Moreover, in communities
of regional provenance more sown species and fewer unsown
species grew compared with communities of foreign provenances indicating higher competitive ability against local
weeds of regional plant communities than of foreign ones.
Thus, our results suggest the use of local or regional seed
material as an integral part of grassland restoration and management to suppress weeds and reduce proportions of bare
ground. Although the proportion of sown and unsown species
changed, aboveground biomass production for both groups
combined did not differ between regional and foreign plots.
Geographic and climatic variation among
provenances
With increasing climatic mismatch between experimental
site and region of seed origin the plant communities performed more poorly. Although we did not measure plant
performance directly, but used cover by each species as
a surrogate, the observed pattern was consistent with the
expectation of climatic adaptation (Becker et al. 2006). For
example, plants adapted to longer growing seasons, such as
those of Swiss provenance, performed more poorly than the
regional community in our experimental field site adapted
to a relatively short growing season. Thus, seeds from a
warmer climate might not be worse per se, but might still
perform worse in a colder area due to maladaptation, which
might involve the expression of metabolic or physiological
activity suboptimal in new conditions (McKay et al. 2001;
Cavender-Bares 2007). Plant performance can decrease with
increasing geographic distance between experimental site
and site of origin (Joshi et al. 2001; Becker et al. 2006),
but in our experiment, plant cover decreased with increasing geographic distance just in the first year weakly and the
effect did not withstand the statistical adjustment. Clearly, not
all climatic or other factors affecting plant growth necessarily need to be correlated with geographic distance, and thus
adaptive differences in plant performance need not always be
best explained by geographic distance (Raabova et al. 2007).
Conclusions
So far, the relevance of using regional seed material for
establishing individual plants and species is acknowledged
in restoration and forestry (Vander Mijnsbrugge et al. 2010).
In addition, our study suggests that regional adaptation matters substantially for community development and ecosystem
properties. As breeding and choice of agriculturally used
plant species so far are focused on single species without considering their interspecific interactions (Lüscher, Connolly,
& Jacquard 1992), we suggest taking the community context
into account for further breeding and species selection. Our
study highlights that regional seed material also improves
ecosystem properties of establishing plant communities as
they are typical for pastures and meadows.
Acknowledgements
We thank Ilka Egerer for suggesting the field site, the
Wachter family for field site access, Ursula Winkler, Antje
Thondorf for lab assistance, many students for field assistance, Alexander Harpke, Sven Pompe, Oliver Schweiger
and the ALARM-Project for climate data, the plant population biology group at the UFZ Halle for discussion, Kathryn
Barto for improving our English and previous reviewers for
comments.
Appendix A. Supplementary data
Supplementary data associated with this article can be
found, in the online version, at http://dx.doi.org/10.1016/j.
baae.2012.07.004.
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