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The Importance of Plant Provenance and Genotypic Diversity of Seed

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The Importance of Plant Provenance and Genotypic Diversity of Seed Published in "Restoration Ecology : doi:10.1111/j.1526-100X.2008.00454.x, 2009" which should be cited to refer to this work. The Importance of Plant Provenance and Genotypic Diversity of Seed Material Used for Ecological Restoration Armin Bischoff,1,2,3 Thomas Steinger,4 and Heinz Mu¨ ller-Scha¨rer1 Abstract the local population. The productivity was greater in pop- The increased translocation of plant species for biodiver- ulations of high genotypic diversity than in those of low sity restoration and habitat creation has provoked a debate diversity, but the effect was only significant in one species. on provenance and genotypic diversity of the used plant Productivity was also more constant among populations material. Nonlocal provenances are often not adapted to of high diversity, reducing the risk of establishment fail- the local environmental conditions, and low population ure. Our results indicate that the choice of an appropriate genotypic diversity may result in genetic bottlenecks ham- provenance and a sufficient genotypic diversity are impor- pering successful establishment. We tested provenance dif- tant issues in ecological restoration. The use of local prov- ferentiation of four plant species used in agri-environment enances does not always guarantee the best performance, schemes to increase biodiversity of agricultural landscapes but a spread of superior alien genotypes can be avoided. A (wildflower strips). Provenances were collected close to sufficient genotypic diversity of the sown plants might be the experimental field and at four further sites of different a biological insurance against fluctuations in ecosystem pro- distances ranging from 120 to 900 km. In two of these cesses increasing the reliability of restoration measures. provenances, different levels of genotypic diversity were simulated by sowing seed from a high and low number of mother plants. We found a large provenance differentia- Key words: competition, diversity–productivity relation, tion in fitness-related traits, particularly in seedling emer- genetic diversity, population differentiation, seed origin, gence. There was no evidence for a general superiority of wildflower strips. Introduction 2006b). However, the required translocation of plant mate- Human impact has reduced species diversity in many eco- rial has provoked a discussion on provenance and diversity systems as a result of habitat damage or even destruction. of the introduced populations (Hamilton 2001; Wilkinson During recent years, interest has increased to restore bio- 2001; Hufford & Mazer 2003). Recent studies have shown http://doc.rero.ch diversity and recreate habitats. Simply stopping negative that the approach has potential risks that may counteract impact and recreating appropriate environmental condi- the desired effects if nonlocal provenances are used and tions are often not sufficient to reestablish species-rich genotypic diversity of the source populations is too low communities of native plants because nearby populations (Keller et al. 2000; Williams 2001; McKay et al. 2005). that are required as propagule sources are not available The risks of introducing nonlocal provenances include any more (Bischoff 2002; Walker et al. 2004). Thus, sow- poor establishment of sown provenances owing to poor ing or planting of desired species has been increasingly adaptation to the prevailing environmental conditions used to improve restoration success. Fields of application (Hamilton 2001; Hufford & Mazer 2003), superior alien range from reintroduction of rare species to sowing spe- genotypes invading and displacing local populations cifically designed seed mixtures for habitat creation (Saltonstall 2002), and hybridization between introduced (Montalvo et al. 1997; Bullock et al. 2001; Bischoff et al. and local populations that may result in outbreeding depression, that is, a reduction of hybrid fitness relative to their parents (Fenster & Galloway 2000; Keller et al. 2000; 1 Department of Biology, Unit of Ecology and Evolution, University of Montalvo & Ellstrand 2001). Theory on local adaptation Fribourg, Chemin du muse´e 10, CH-1700 Fribourg, Switzerland 2 Address correspondence to A. Bischoff, email [email protected] predicts that local genotypes should perform better than 3 Present address: Department of Biological Sciences, National Institute distant provenances when grown at their home site. Such of Horticulture and Landscape Planning (INHP), Agrocampus West, Center of Angers, 2, rue Andre´ Le Noˆ tre, F-49000, Angers, France an adaptive genetic differentiation is well documented in 4 Station de recherche Agroscope Changins-Wa¨denswil ACW, Case postal plant populations (Bradshaw 1984; Linhart & Grant 1996; 1012, CH-1260 Nyon, Switzerland Galloway & Fenster 2000; Etterson 2004). Generally, adaptation to a specific site is expected to decrease with increasing spatial distance from the source population due 1 to a concomitant increase in environmental distance and extremes (Reusch et al. 2005). Due to differences among genetic isolation (Keller & Kollmann 1999; Galloway & genotypes in their resistance to herbivory and abiotic Fenster 2000; Joshi et al. 2001). However, genetic differ- stress, the chance that populations comprise appropriate entiation has also been shown on very small scales or even genotypes increases with genotypic diversity. Hence, vari- within populations (Waser & Price 1985; Knight & Miller ation in fitness among diverse populations is supposed to 2004; Lenssen et al. 2004; Bischoff et al. 2006a), reducing be lower than in nondiverse populations, and the risk of the correlation between fitness and spatial distance of failure in establishment of single population may be a provenance. smaller. In our study, we tested for effects of genotypic In the present study, we analyzed effects of seed prove- diversity on mean population productivity and among nance in four plant species of an established wildflower population variability in two plant species. We manipu- seed mixture used to create wildflower strips in agricul- lated diversity levels by varying the number of seed fami- tural landscapes. Such wildflower strips have been imple- lies sown into experimental plots. mented in Switzerland as ecological compensation areas We tested both provenance and diversity effects in to restore species richness and cover currently 2,400 ha on plants growing with and without surrounding vegetation 2,600 different farms (Swiss Federal Office for Agriculture in order to examine whether they depend on the competi- 2005). We tested five provenances of the four species in tion regime. Most earlier studies have been conducted a field experiment, that is, one local provenance and four under noncompetitive conditions, assuming that the fit- provenances collected at different geographical and envi- ness ranking of the populations remains the same under ronmental distances to the field site. Our aim was to ana- natural competitive conditions (Bennington & McGraw lyze whether there is significant population differentiation 1995; Keller & Kollmann 1999; Joshi et al. 2001). How- between provenances and whether performance is nega- ever, such a correlation may be weak or absent, and other tively correlated with distance to the collection sites. Envi- experiments have illustrated the importance of the sur- ronmental maternal effects may contribute to population rounding vegetation as a potential driver of local adapta- differentiation in plants (Roach & Wulff 1987; Donohue & tion (Prati & Schmid 2000; Ehlers & Thompson 2004; Schmitt 1998). In a previous study on the same five popu- Bischoff et al. 2006a). lations of the test species, seed weight contributed little to We addressed the following specific research questions: among-provenance variation in germination and seedling (1) How strong is population differentiation among five performance, suggesting that maternal effects were rela- different European provenances in each of four spe- tively small (Bischoff et al. 2006b). To further check this cies and do the local provenances perform the best? assumption, we additionally grew F1 seeds of one test spe- (2) What is the effect of genotypic diversity on mean popu- cies produced under homogeneous conditions and com- lation productivity and among population variability? pared the provenance ranking in the F1 generation with that of the parental generation. Intraspecific diversity of the introduced plant material is a second issue that may be important for ecological res- Methods toration, but few empirical studies have addressed this http://doc.rero.ch issue. If seeds are collected from a limited number of sour- Study Species, Provenances, and Sampling ces, the level of genetic diversity is low and genetic bottle- We selected four species of a standard seed mixture used necks may occur in the restored populations (Hufford & to establish wildflower strips as ecological compensation Mazer 2003). Procaccini and Piazzi (2001) observed that areas in Swiss agricultural landscapes. They represent dif- genotypic diversity of the founders may have stronger ferent stages of succession from early to late: the annual effects on the fitness of a population than the plant prove- Legousia speculum-veneris (L.) Chaix (Primulaceae), the nance. Recent studies have shown that intraspecific diver- biennial Echium vulgare L. (Boraginaceae), the short- sity, like species diversity, may be positively correlated lived biennial or perennial Cichorium intybus L. (Astera- with productivity and ecosystem functioning (Reusch
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