Professional Documents
Culture Documents
TMP A569
TMP A569
Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec
Original article
a r t i c l e i n f o a b s t r a c t
Article history: Several longer-term assembly studies on ex-arable land have found that species that arrive first at a
Received 29 April 2013 disturbed site can play a key role in the further development of the community and that this priority
Accepted 16 September 2013 effect influences aboveground productivity, species diversity and stability of the grassland communities
Available online
that develop. Restoration of nutrient poor, species rich grasslands is often limited by seed dispersal as
well as the accessibility of suitable microsites for establishment. Sowing species (i.e. creating priority
Keywords:
effects for further assembly) may help overcome such dispersal barriers, but the potential of using
Community assembly
priority effects for restoration has not been tested in this type of dry grassland. We tested the hypothesis
Biodiversity
Restoration
that sowing two different seed mixtures used for dry acidic grassland restoration onto a sandy substrate
Filter theory (which formed an equivalent to a primary succession) would create priority effects, and that these
Initial sowing priority effects would be sustained over a number of years. We followed community assembly and
Invasion measured aboveground productivity for four years after sowing. We found that priority effects caused by
Microsite limitation sowing of differently diverse mixtures did also occur in dry acidic grassland habitat, but that how
persistent they were over time depended on the response variable considered. Priority effects on species
number were not as strong as found in previous ex-arable land studies, whereas priority effects for
aboveground productivity were still visible after 4 years. In addition, functional composition of the
community still reflected the composition of the seed mixtures 4 years later. Our results suggest that
priority effects can occur in nutrient-poor dry acidic grassland but in contrast to more nutrient-rich sites
the breadth of responses affected may not be as wide.
Ó 2013 Elsevier Masson SAS. All rights reserved.
1146-609X/$ e see front matter Ó 2013 Elsevier Masson SAS. All rights reserved.
http://dx.doi.org/10.1016/j.actao.2013.09.004
C. Plückers et al. / Acta Oecologica 53 (2013) 110e116 111
Bezemer and van der Putten (2007) performed an experiment grassland created by sowing two different seed mixtures at time
sowing either zero, four or fifteen species of plants onto ex-arable zero onto sandy substrate which formed an equivalent to a primary
land then followed the dynamics of the system in terms of spe- succession and asked the following two questions:
cies turnover, productivity, temporal stability and diversity. In a
grassland restoration context, Bullock et al. (2001, 2007) sowed 1) Does sowing two different seed mixtures produce priority ef-
either low or high diversity mixtures using plant mixtures typical fects in dry grassland, and how sustainable are they over time?
for UK grassland restoration on a whole series of ex-arable sites and 2) If there are priority effects which traits, processes or charac-
over a long period of time. Both studies found long-lasting effects teristics of the ecosystem do they relate to most?
(hereafter called priority effects) of initial sowing of seeds on
further assembly, in particular when more diverse seed mixtures In order to test this, we followed changes in different traits of
were sown. Flombaum and Sala (2008) removed species to create a the community over 4 years: at community level total species
plant species diversity gradient in the Patagonian steppe and found number (SN), community cover and aboveground peak biomass as
that aboveground net primary production increased with the a surrogate of productivity. We also assessed differences between
number of plant species. responses of target (desired sown) and non-target species (in-
The restoration of species-rich communities is becoming a vaders). At functional group level total species number, community
major tool to counteract biodiversity loss but it can also have cover and total aboveground productivity were split into functional
positive effects on some ecosystem functions, for example groups (grasses, legumes, non-legume forbs).
increasing biomass production and hence nutrient cycling and
reducing erosion (UNEP Nagoya Protocol, 2011). In a world expe- 2. Materials and methods
riencing increasing global change, where historical reference sys-
tems often no longer exist, ecological restoration is adapting by 2.1. Experimental design
focusing as much on ecosystem functioning as on species compo-
sition when setting restoration goals (Choi et al., 2008. Hobbs et al., In autumn 2007 we established a grassland assembly experi-
2009). ment, the Habitat Garden, with two different grassland habitats
(dry acidic and mesic grassland). The experimental field plots are
1.1. Using priority effects for restoration on the campus of the Forschungszentrum Jülich, located in Jülich,
West Germany (6 220 000 E, 50 560 000 N), and consist of 12 plots
Priority effects occur when species that arrive first in an (randomized, each 2 2 m in size and separated by 50 cm paths
ecosystem significantly affect the further development of the sown with a non-clonal grass species). The dry acidic grassland
community and thereby strongly influence community compo- plots (n ¼ 6) were sown with two different diversity mixtures
sition (Facelli and Facelli, 1993; Fukami et al., 2005). Priority ef- (whereas the mesic plots were sown with the same diversity of
fects can lead to lasting differences in species or functional group species at the start for removal experiments (not considered here,
dominance, and hence can potentially drive ecosystem properties see also Plückers et al. 2013)).
and functioning. Priority effects can occur on timescales from The dry acidic grassland part of the experiment was designed to
days to years and can be linked to differences in the arrival time follow possible priority effects of sowing differently diverse mix-
of a species at a site but also to their success in establishing tures on community assembly over time on a sandy substrate
themselves in the community. Priority effects can short-term or which thus formed an equivalent to a primary succession. For each
lead to alternative stable states in vegetation (Grman and Suding, dry acidic grassland plot the original soil was removed by digging
2010). As such, restoration projects are often contingent upon out the soil to a depth of 40 cm, and a geomembrane permeable to
priority effects in that they can alter vegetation trajectories as water and nutrient laid down to avoid root input from plants
well as impede restoration success in some cases. The mecha- growing outside the plots and to remove any seed bank. The sandy
nisms underlying the priority effects or the timescale upon which substrate, which consisted of sand (grain size 0.7e1.4 mm) mixed
they operate are rarely addressed however (Grman and Suding, with one tenth potting soil (with very low nutrient and availability)
2010). was then filled into the prepared holes.
Most of the evidence for priority effects comes from grassland We sowed two differently diverse seed mixtures at a density of
systems with intermediate soil nutrient availability (see Bezemer 3 g/m2 using typical restoration mixes used to establish dry acidic
and van der Putten, 2007; Bullock et al., 2001, 2007). We do not grasslands in Germany (Rieger Hofmann GmbH Blaufelden, Ger-
know whether priority effects of sowing different mixtures can play many), in December 2007. There were two diversity treatments
a role in more nutrient-poor dry grasslands. Foster and Dickson (n ¼ 3 per sowing treatment): S2 consisted of 2 grass and 25 forbs
(2004) hypothesise that systems with higher resource availability (one of which was a legume), and S7 consisted of 7 grass and 32
have more available niches but these niches are usually easily filled forb species (four of which were legumes). Species within the lower
(packed) with species and this creates more neighborhood diversity S2 treatment formed a subset within the mixtures sown
competition. In contrast in systems with more limited availability for treatment S7: e.g. the 2 grass species in S2 were also part of the
of resources such as soil nutrients, species attempting to establish S7 mixtures, the one legume species sown in S2 was also a subset of
themselves generally may find more open niche space but mainly S7, Both mixture treatments had 12 non-legume forb species in
face microsite limitation posed by abiotic conditions. Given this, common, whereas S7 has 16 additional different non-legume forb
one might expect that sowing differently diverse dry acidic grass- species and S2 has 12 additional different non-legume forb species.
land mixtures on sandy substrates (which form an equivalent to a These seed mixtures were chosen, to ensure the study had some
primary succession) would overcome dispersal barriers but that potential for regional restoration application in the future, such
microsite and nutrient limitations may deter establishment of that we chose seed mixtures typically used by restoration practi-
target species more than on a more mesic substrate. The timescale tioners and land managers in central Germany, provided by the
upon which priority effects may operate in dry acidic grasslands wild seed company Rieger Hofmann GmbH. One quarter of each
may therefore be shorter than for more mesic sites. plot was not sown and kept as a control non-sown subplot. The
Our field study aimed to test the strength of priority effects (in experiment was fenced off to reduce confounding factors such as
terms of detection of priority effects over time) in a dry acidic grazing by deer or wild boar. The plots were mown once a year in
112 C. Plückers et al. / Acta Oecologica 53 (2013) 110e116
Fig. 1. The development of species number and cover at the community and functional group level. Values are means (þone standard error of the mean) A and C: Community level:
note that plant cover for a plot can be higher than 100%, because of 3-D aspects of plant community canopies. B. and D. Functional group level: non-legume forb species (bottom
section), legume species (middle section) and grasses (top section).
significant year effect), but the detailed development of both (Fig. 2B, Table 1 significant treatment effect). The relative above-
treatments was different over time, which was confirmed by a ground biomass production of grasses, forbs and legumes did not
significant interaction effect between treatment and year effects differ significantly between 2010 and 2011 (when measured). Total
(Fig. 2A, Table 1). aboveground biomass production increased in 2011 in the S7 plots,
The S7 plots had a higher total aboveground biomass production due to a relative increase in legume biomass (especially of Lotus
in all years (Fig. 2A, Table 1 significant treatment effect). Total corniculatus). The high variability of the error bars for 2011 biomass
aboveground biomass of legumes was much higher in the S7 plots can be mainly attributed to the presence of one woody legume
than in the S2 plots, whereas forbs dominated more in S2 plots shrub in one plot (Genista tinctoria).
Fig. 2. The development of aboveground biomass at the community and functional group level. Values are means (þone standard error of the mean) A. Community level B.
Functional group level (as in Fig. 3). The total aboveground biomass values of the community level and the sum of the functional group level is not the same because of trans-
formation procedures.
114 C. Plückers et al. / Acta Oecologica 53 (2013) 110e116
Invasion of species increased over time but species richness was Sowing treatment % Establishment of target species
not significantly different between treatments (Table 1 see Fig. 3A 2008 2011
total number of species). Species number and cover of non-target S2 28.39 2.5 40.74 3.7
species (non-sown species) and target species (sown species) of S7 31.62 5.2 36.75 4.3
both sowing treatments increased over time (Fig. 3A, Table 1 sig-
nificant year effect). There was a significant effect of the sowing
treatments on the cover of target species (Table 1 significant sow- species such as thistle (Cirsium, Equisetum, Rumex or Polygonum sp).
ing treatment), but no significant effect of the sowing treatments Non-target dry acidic grassland species from other sandy plots
on the number of non-target species and their cover (see Table 1 no formed an intermediate group of species that successfully managed
treatment effect overall years). The ratio of target to non-target to invade at a later time point after sowing. This is as one might
species number significantly decreased over time as newcomers expect, given the number of sown species that the treatments
arrived (Table 1 significant year effect, Table 2 for changes in per- shared in common, such that quite a few of the sown dry acidic
centage target species). At the beginning 27 species were sown in grassland species in S2 treatments were a subset of the species in
the S2 plots and 39 species in the S7 plots. Only 30e40% of the S7. In detail, there were 12 “new” non-sown target species that
sown species had established after 4 years in 2011 (Table 2), with could potentially invade from S2 to S7, versus 24 options from S7 to
both sowing treatments having nearly 30% target species in the first S2. Our results follow the expected relative success of target species
year (2008) but had slightly higher establishment in the S2 plots (those sown on a treatment) with high invasion from S7 to S2 than
than S7 by 2011 (Table 2). Overall, S7 plots had a higher estab- the other way. Over the 4 years, the sowing treatment had a sig-
lishment of target species (relative to non-target species) than S2 nificant effect on assembly. The control plots had significantly less
plots (Table 2), but the proportion of established target species in total species number, total cover and total aboveground biomass
the S2 plots significantly increased from 2008 to 2011 (t-test production over the whole time than the sown plots (data not
p ¼ 0.050, data not shown), whereas this was not the case in S7. The shown).
non-target species that managed to establish did not mainly derive
from the surrounding mesic grassland plots, but were ruderal 3.3. Did sowing differently diverse mixtures affect soil conditions?
4. Discussion
dependent on which variable one measures. It would be interesting establish by 2011 in both sowing treatments. Our data fit in well
to follow more ecosystem functions than just aboveground pro- with results from central European grassland restoration projects
ductivity in further studies, as well as typical vegetation measures. (Kiehl et al. 2010) where 4 years after sowing (also at a density of
One possible reason for the milder priority effects we found on dry 3 g/m2) around 30e80% of target species had managed to establish,
acidic grassland compared to mesic conditions, could be that in and the lower establishment rates were mainly on low-nutrient
communities developing on very sandy substrates, the key limita- sites. This suggest that in our study microsite limitation was play-
tions to establishment (apart from dispersal) are abiotic in nature ing a role and affected species richness of the vegetation, even if we
(e.g. microsite limitation) and less driven by direct interactions did not directly measure it. Kiehl et al. (2010) assessed various
between plant species already present (see Fig. 4). As Foster and techniques for introducing species to a site during restoration in
Dickson (2004) highlight in their conceptual model of how inva- Central Europe and found that although sowing could overcome
sion is modulated by available nutrient resources for plants: sys- dispersal limitation, the long-term success of restoration also
tems with higher resource availability experience more niche depended very much on the availability of appropriate abiotic
packing and hence more competition than systems with lower conditions including establishment microsites.
resource availability. Invading species experience lower microsite In classical biodiversity-ecosystem functioning experiments
limitation however, since abiotic conditions are more benign. (where natural assembly is not allowed) more diverse communities
Sowing more diverse seed mixtures should initially increase the are generally more resistant to invasion (Roscher et al, 2009).
niche packing speed since it removes dispersal limitation, however Huang et al. (2013) found in a prairie biodiversity experiment
at later successional stages the area with less seeds sown should where weeding ceased after 3 years, that the positive relationship
allow more new invaders to establish than the high diversity site. In between diversity and productivity persisted even after cessation
contrast in more nutrient-limited systems, species attempting to of weeding. In our study invasion pressure did increase total species
establish should generally find more open niche space but mainly number but the lack of significant difference in species number
face microsite limitations posed by abiotic conditions, as we found between the sowing treatments (except in year 1) suggests that
in this study. Sowing more diverse mixtures should initially invasion resistance may have been similar between treatments
decrease dispersal limitation and increase establishment of target (Fig. 1). Overall, S7 plots had a higher establishment of target spe-
species. In later succession (in contrast to the higher resource cies (relative to non-target species) as well as higher soil C content
scenario) a higher proportion of the established species will be new and a trend to higher N content than S2 plots (Table 1). The pro-
invaders (non-target) since a lower proportion of the sown species portion of established target species in the S2 plots significantly
will be able to establish due to microsite limitation. During later increased from 2008 to 2011, whereas this was not the case in S7,
succession, facilitation by nurse plants may help new invaders to despite soil C remaining higher in the S7 treatment. So S7 started
establish, and the potential for this may increase in more diverse out with higher establishment success for target species but S2
sites. caught up over time (Table 2). This is mirrored in the cover data
By sowing two differently diverse seed mixtures on a sandy where S2 started out having lower community cover than S7 but
substrate we influenced the availability of propagules and reduced become significantly higher than S7 by 2010 and 2011 (see Fig. 1C,
the dispersal limitation typical of such nutrient-poor grassland see 3B for target/non target species cover). This shows that the two
communities in this early stage of assembly. We made species treatments may have had similar invasion of species (in terms of
available through sowing and gave these species the chance to numbers) but that the abundance of the species differed.
establish themselves first and thus cause priority effects in further It seems that time plays a crucial role in restoration success for
assembly. Considered within filter theory (Kelt et al., 1995): to establishing desired target species in species-rich grasslands. Baasch
establish themselves, our sown seeds (target species) had to pass et al. (2012) evaluated restoration experiments in ex-mining sites on
through the mesh of the abiotic filter (since the sandy substrate sandy soils in eastern Germany and found that the species-rich
formed an equivalent to a primary succession with extremely low grasslands established after hay transfer or sowing were highly
nutrient and water holding capacity). The sown target species did resistant to invasion of ruderal species (despite hay transfer
not however have to overcome a biotic filter resulting from plant methods not only having positive effects on establishment). After 9
species already present in the habitat at the moment of their years, however, there was no difference between treatments in
arrival. At this time point we expect that microsite limitation will terms of total vegetation cover, species richness and the number of
have affected the germination and establishment success of the target species. In our study we found the same effect after only 4
seeds the most. Our establishment data (Tables 2 and 3) back this years: there was no difference between sowing treatments in terms
up, with around only 30% of the sown target species managing to of total species number and number of target species, although
establish in the first growing season, and around 40% managing to productivity, community cover and functional composition did vary.
Focusing on the different functional groups of the species sown
(e.g. legumes, non-legume forbs and grasses) our study found that
the functional group composition sown was still detectible in the
vegetation 4 years after sowing. This is an interesting effect, even if
we cannot separate effects of the species richness from the func-
tional richness of the seed mixtures.
This detectable priority effect after 4 years was valid for cover
and biomass of forbs and legumes, and richness of legumes only
(Table 1) but not for grasses. While one can see in our data that the
higher proportion of legumes sown was reflected in higher cover
Fig. 4. Conceptual model of filter theory of community assembly based on Kelt et al., and biomass of legumes over time, this effect was not found for the
1995 (modified from Hobbs and Norton, 2004) and adapted to include facilitation as forbs. For the forbs, the S2 treatment that started out with less forb
well as competition as part of the mechanisms behind the biotic filter. The dotted species, had an as high proportion of forbs in the community after 4
arrow highlights when facilitation can help a species manage to establish, e.g. if a
nurse plant provides a microsite or additional nitrogen during germination and early
years as the S7 treatment. The particularly strong priority effect of
growth. The strongly dotted arrow indicates negative competitive interactions sowing legumes on legume composition may be related to the
potentially blocking a species from managing to establish. finding that legumes established quickly and well. They seemed to
116 C. Plückers et al. / Acta Oecologica 53 (2013) 110e116
be well adapted to the conditions on site from the start and Bullock, J.M., Pywell, R.F., Burke, M.J.W., Walker, K.J., 2001. Restoration of biodi-
versity enhances agricultural production. Ecol. Lett. 4, 185e189.
established well in S7, whereas in S2 the one legume species did not
Bullock, J.M., Pywell, R.F., Walker, K.J., 2007. Long-term enhancement of agricultural
establish well, allowing the non-legume forbs to become dominant. production by restoration of biodiversity. J. App. Ecol. 44, 6e12.
We know from many greenhouse and field studies with legumes Choi, Y.D., Temperton, V.M., Allen, E.B., Halassy, M., Hobbs, R.J., Grootjans, A.P.,
interacting with other functional groups, that legumes tend to be Naeth, M.A., Torok, K., 2008. Ecological restoration for future sustainability in a
changing environment. Ecoscience 15, 53e64.
competitive across a range of abiotic conditions (Temperton et al., Facelli, J.M., Facelli, E., 1993. Interactions after death: plant litter controls priority
2007; non-published data). effects in a successional plant community. Oecologia 95, 277e282.
If in follow-up experiments the functional composition of the Flombaum, P., Sala, O.E., 2008. Higher effect of plant species diversity on produc-
tivity in natural than artificial ecosystems. Proc. Natl. Acad. Sci. U S A 105,
mixture were found to be more important than the species rich- 6087e6090.
ness, one could perhaps use priority effects of initial sowing Foster, B.L., Dickson, T.L., 2004. Grassland diversity and productivity: the interplay
composition to direct the functional composition of the community of resource availability and propagule pools. Ecology 85, 1541e1547.
Fukami, T., Bezemer, T.M., Mortimer, S.R., van der Putten, W.H., 2005. Species
as well as total aboveground biomass and cover. This is turn could divergence and trait convergence in experimental plant community assembly.
potentially have positive effects on nutrient cycling and carbon Ecol. Lett. 8, 1283e1290.
sequestration in mesic grasslands (sensu Steinbeiss et al. 2008; Grman, E., Suding, K.N., 2010. Within-year soil legacies contribute to strong priority
effects of exotics on native California grassland communities. Restor. Ecol. 18,
Oelmann et al. 2011). This would need to be tested in separate 664e670.
experiments before being applicable to restoration since seed Hobbs, R.J., Norton, D.A., 2004. Ecological filters, thresholds, and gradients in
mixtures are rarely separated into these groups, but just sown as resistance to ecosystem reassembly. assembly rules and restoration ecology:
bridging the gap between theory and practice. In: Temperton, V.M., Hobbs, R.J.,
higher or lower diversity.
Nuttle, T., Halle, S. (Eds.). Island Press, Washington DC, USA, pp. 72e95.
Hobbs, R.J., Higgs, E.S., Harris, J.A., 2009. Novel ecosystems: implications for con-
5. Conclusions servation and restoration. Trends Ecol. Evol. 24, 599e605.
Huang, Y., Martin, L.M., Isbell, F.I., Wilsey, B.J., 2013. Is community persistence
related to diversity? A test with prairie species in a long-term experiment. Basic
Overall, our study aimed to test whether priority effects of Appl. Ecol. 14, 199e207.
sowing differently diverse seed mixtures play a role in dry acidic Jentsch, A., Friedrich, S., Steinlein, T., Beyschlag, W., Nezadal, W., 2009. Assessing
grasslands, and we found that they do, but how sustained they conservation action for substitution of missing dynamics on former military
training areas in central Europe. Restor. Ecol. 17, 107e116.
were depended on the response variable measured. Our study Kelt, D.A., Taper, M.L., Meserve, P.L., 1995. Assessing the impact of competition on
found relatively low establishment success of target species, but the community assembly: a case study using small mammals. Ecology 76, 1283e
results are in line with results from low-nutrient grassland resto- 1296.
Kiehl, K., Kirmer, A., Donath, T.W., Rasran, L., Hölzel, N., 2010. Species introduction
ration, suggesting that microsite limitation and related filtering in restoration projects e evaluation of different techniques for the establish-
effects of severe abiotic environments may be the strongest driving ment of semi-natural grasslands in Central and Northwestern Europe. Basic
factors in assembly of dry acidic grassland. Additional reduction of Appl. Ecol. 11, 285e299.
Kirmer, A., Baasch, A., Tischew, S., 2012. Sowing of low and high diversity seed
microsite limitation via planting out nurse plants to facilitate mixtures in ecological restoration of surface mined-land. App. Veg. Sci. 15, 198e
establishment in such harsh conditions or including an interme- 201.
diately severe disturbance regime (as in Jentsch et al. 2009) may be Londo, G., 1976. The decimal scale for relevees of permanent quadrats. Plant Ecol.
33, 61e64.
as important to improving dry grassland restoration success as
Milchunas, D.G., Lauenroth, W.K., 1993. Quantitative effects of grazing on vegetation
sowing therefore. and soils over a global range of environments. Ecol. Mono 63, 327e366.
Nagoya Protocol, 2011. Nagoya Protocol on Access to Genetic Resources and the Fair
Acknowledgments and Equitable Sharing of Benefits Arising from Their Utilization to the
Convention on Biological Diversity. Secretariat of the Convention on Biological
Diversity, Montreal, Convention on Biological Diversity United Nations.
The Habitat Garden Experiment was made possible by Oelmann, Y., Buchmann, N., Gleixner, G., Habekost, M., Roscher, C., Rosenkranz, S.,
Foschungszentrum Jülich funding for the group of Vicky Temper- Schulze, E.-D., Steinbeiss, S., Temperton, V.M., Weigelt, A., Weisser, W.W.,
Wilcke, W., 2011. Plant diversity effects on aboveground and belowground N
ton. We thank all people who helped with field work especially pools in temperate grassland ecosystems: development in the first 5 years after
Edelgard Schölgens and Marlene Müller, and the central chemical establishment. Global. Biogeochem. Cycles 25, GB2014.
laboratory of the Forschungszentrum Jülich (ZEA) for help in Peet, R.K., Glenn-Lewin, D.C., Wolf, J.W., 1983. Prediction of man’s impact on plant
species diversity. In: Holzner, W., Werger, M.J.A., Ikusima, I. (Eds.), Man’s Impact
analyzing soil samples. We thank Marcus Wagner and James on Vegetation. Junk Publishers, den Haag, NL, pp. 41e54.
Bullock from CEH in England for constructive discussions of the Plückers, C., Temperton, V.M., Erler, A., Putz, A., Scharr, H., Rascher, U., 2013. Moving
data. towards measuring multifunctionality in ecosystems: FieldScreen e a mobile
positioning system for non-invasive measurement of plant traits in field ex-
periments. Nova Acta Leopold. 391, 221e237.
References Roscher, C., Temperton, V.M., Buchmann, N., Schulze, E.-D., 2009. Community as-
sembly and biomass production in regularly and never weeded experimental
Baasch, A., Kirmer, A., Tischew, S., 2012. Nine years of vegetation development in a grasslands. Acta Oecol. 35, 206e217.
postmining site: effects of spontaneous and assisted site recovery. J. App. Ecol. Steinbeiss, S., Beßler, H., Engels, C., Temperton, V.M., Buchmann, N., Roscher, C.,
49, 251e260. Kreutziger, Y., Baade, J., Habekost, M., Gleixner, G., 2008. Plant diversity posi-
Bakker, J.P., Berendse, F., 1999. Constraints in the restoration of ecological diversity tively affects short-term soil carbon storage in experimental grasslands. Glob.
in grassland and heathland communities. Trends Ecol. Evol. 14, 63e68. Chang. Biol. 14, 2937e2949.
Balvanera, P., Pfisterer, A.B., Buchmann, N., He, J.-S., Nakashizuka, T., Raffaelli, D., Temperton, V.M., Mwangi, P.N., Scherer-Lorenzen, M., Schmid, B., Buchmann, N.,
Schmid, B., 2006. Quantifying the evidence for biodiversity effects on ecosystem 2007. Positive interactions between nitrogen-fixing legumes and four different
functioning and services. Ecol. Lett. 9, 1146e1156. neighbouring species in a biodiversity experiment. Oecologia 151, 190e205.
Bezemer, T.M., van der Putten, W.H., 2007. Ecology: diversity and stability in plant Tylianakis, J.M., Didham, R.K., Bascompte, J., Wardle, D.A., 2008. Global change and
communities. Nature 446, E6eE7. species interactions in terrestrial ecosystems. Ecol. Lett. 11, 1351e1363.