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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.2012.07.004 552 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). 553 554 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. 556 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 558 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. References Becker, U., Colling, G., Dostal, P., Jakobsson, A., & Matthies, D. (2006). Local adaptation in the monocarpic perennial Carlina vulgaris at different spatial scales across Europe. Oecologia, 150, 506–518. 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