Summary 1. There are few studies of the performance of species in restored vegetation communities. Here we report the results of a meta-analysis of 25 experiments concerned with species-rich grassland restoration on ex-arable land and agriculturally improved grasslands situated at a wide range of locations throughout lowland Britain. Differences in species’ performance were related to 38 physiological and morphological traits. 2. An experiment-adjusted performance index was calculated for each of the 58 species (13 grasses and 45 forbs). The performance index was calculated for the first 4 years after establishment together with a temporal trend. 3. Individual species showed large differences in performance indices. However, grasses consistently out-performed forbs. 4. We examined the linkage between species’ performance and traits according to four non-exclusive hypotheses. The ability to establish and persist in restored vegetation communities requires: (H1) good gap colonization ability; (H2) strong competitive capability; and (H3) ability to undergo vegetative regeneration. (H4) Successful species are generalists associated with fertile habitats. 5. Trait analyses supported all four hypotheses. Within the forbs, good establishment in the first year was linked to traits determining colonization ability: ruderality, percentage germination of seeds and autumn germination. However, traits linked to competitive ability, vegetative growth and seed bank persistence became increasingly important determinants of success with time. Species with generalist habitat requirements, and especially those associated with fertile soils, performed increasingly well with time. This reflects the development of a closed vegetation in which the ability to grow vigorously and out-compete other established plants is important. 6. Stress-tolerators, habitat specialists and species of infertile habitats performed badly. This may reflect high residual fertility in restored grasslands and particular niche requirements of these species. This may be a problem as grassland restoration often targets communities characterized by species with these traits and many are food plants of invertebrates of conservation value. 7. There were few significant correlations between the performance of the grasses and traits reflecting their overall good performance in comparison with the forbs. 8. This study has important implications for practical restoration programmes and policies. Efficiency might be increased by introducing only species with good performance, but this would lead to uniformity among restored grasslands and would diminish the benefits of habitat restoration for national and regional biodiversity. 9. Synthesis and applications. Future work should focus on practical methods to increase the successful establishment of the poor performing but desirable species, by (i) targeting restoration to low fertility soils, (ii) changing the abiotic environment or (iii) the ‘phased

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1046/j.1365-2664.2003.00762.x

Plant traits as predictors of performance in ecological restoration

frequently with a growth of herbs in its understorey which contrasts with the surrounding areas by its higher biomass and diversity. We examined the relationships between the shrub and the herbs underneath along a gradient of shrub age. A total of fifty individuals were selected to fit five age classes and shrub characteristics, soil properties and flora under the canopy were examined along the estimated chronosequence. All shrub size variables increased with time, as did the amount of nutrients stored by the shrub, but differences were often significant only between the three oldest classes. Concentration of nitrogen and phosphorus in photosynthetic stems remained constant, but nutrient pools in stem biomass increased with time. The shrubs changed the soil environment under their canopies with age by ameliorating soil texture, nutrient content and capacitance of water. The improvement was most pronounced between the two oldest classes (IV and V), and was probably due to the high biomass of perennial species in the understorey which increased the production of litter, and the interception of wind-blown dust. Plant diversity in the understorey increased with shrub age, likely due to a greater heterogeneity under larger canopies. Drought-resistant species, typical of the open areas between shrubs, were displaced from the centre of the understorey by taller, more mesic species over the age gradient. Retama sphaerocarpa shrubs benefited from the increase of resources in the understorey and showed a higher reproductive output with shrub age, but decreased at the beginning of senescence. The indirect interactions between R. sphaerocarpa and its understorey herbs could be considered as a two-way facilitation in which both partners benefit from their association.

http://www.jstor.org/stable/pdfplus/3546339.pdf

Facilitation and succession under the canopy of a leguminous shrub, Retama sphaerocarpa, in a semi-arid environment in south-east Spain

The orchid Dactylorhiza sambucina shows a stable and dramatic flower-color polymorphism, with both yellow- and purple-flowered individuals present in natural populations throughout the range of the species in Europe. The evolutionary significance of flower-color polymorphisms found in many rewardless orchid species has been discussed at length, but the mechanisms responsible for their maintenance remain unclear. Laboratory experiments have suggested that behavioral responses by pollinators to lack of reward availability might result in a reproductive advantage for rare-color morphs. Consequently, we performed an experiment varying the relative frequency of the two color morphs of D. sambucina to test whether rare morph advantage acted in the natural habitat of the species. We show here clear evidence from this manipulative experiment that rare-color morphs have reproductive advantage through male and female components. This is the first demonstration, to our knowledge, that negative frequency-dependent selection through pollinator preference for rare morphs can cause the maintenance of a flower-color polymorphism.

https://www.pnas.org/content/pnas/98/11/6253.full.pdf

Negative frequency-dependent selection maintains a dramatic flower color polymorphism in the rewardless orchid Dactylorhiza sambucina (L.) Soo.

Summary 1 Species attributes have been used to explain invasion patterns assuming the prevalence of biological mechanisms, although this approach often suffers several methodological and conceptual limitations, such as local idiosyncrasies, differences among habitats, phylogenetic constraints and insufficient sample size. 2 The relative importance of 15 species traits for explaining the abundance over 350 naturalized alien plant species was assessed across five Mediterranean islands (Corsica, Crete, Majorca, Malta and Sardinia). A comparative analysis accounting for phylogeny was used to examine variation in semi-quantitative estimates of species abundance in comparable habitats across the five island floras. 3 Species were divided into those with affinity for semi-natural, agricultural and ruderal habitats. Both vegetative and reproductive attributes were evaluated for individual islands and averaged across all islands. 4 Vegetative propagation, large leaf size, summer flowering, long flowering period and dispersal by wind or vertebrates were positively associated with average alien abundance across all five islands. Fewer significant trends were found in island-specific patterns. 5 The relative importance of a few reproductive traits is reflected in over-representation of Caryophyllales, Asterales and Poales (late flowering, large seed size and anemochory). Although significant covariation in traits was found there was no evidence for welldefined invasive syndromes. 6 Succulence was important in ruderal habitats, long flowering period in agricultural habitats and vertebrate seed dispersal in semi-natural habitats, suggesting that empty niches, avoidance of competitors and exploitation of mutualists, respectively, are important. 7 The study highlights the importance of estimating invasion success across a wide region, but analyses of specific invasion stages are also needed. Reproductive traits, which may be more relevant for long-distance colonization, and vegetative traits, which determine local dominance and persistence, were, nevertheless, both related to abundance within islands.

Species attributes and invasion success by alien plants on Mediterranean islands

Rapid anthropogenic climate change is already affecting species distributions and ecosystem functioning worldwide. We applied niche-based models to analyse the impact of climate change on tree species and functional diversity in Europe. Present-day climate was used to predict the distributions of 122 tree species from different functional types (FT). We then explored projections of future distributions under one climate scenario for 2080, considering two alternative dispersal assumptions: no dispersal and unlimited dispersal. The species-rich broadleaved deciduous group appeared to play a key role in the future of different European regions. Temperate areas were projected to lose both species richness and functional diversity due to the loss of broadleaved deciduous trees. These were projected to migrate to boreal forests, thereby increasing their species richness and functional diversity. Atlantic areas provided an intermediate case, with a predicted reduction in the numbers of species and occasional predicted gains in functional diversity. This resulted from a loss in species within the broadleaved deciduous FT, but overall maintenance of the group. Our results illustrate the fact that both species-specific predictions and functional patterns should be examined separately in order to assess the impacts of climate change on biodiversity and gain insights into future ecosystem functioning.

Using niche-based modelling to assess the impact of climate change on tree functional diversity in Europe

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Flora Europaea Vol. 4, edited by T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters, and D. A. Webb. Cambridge University Press, £25.

A. H. Fitter

Papers

Flora Europaea Vol. 4, edited by T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters, and D. A. Webb. Cambridge University Press, £25.