Plant reproductive strategies vary under low and high pollinator densities

Long‐term variation in the population density of honey bees Apis mellifera across landscapes has been shown to correlate with variation in the floral traits of plant populations in these landscapes, suggesting that variations in pollinator population density and foraging rates can drive floral trait evolution of their host plants. However, it remained to be determined whether this variation in plant traits is associated with adaptive variation in plant reproductive strategies under conditions of high and low pollinator densities. Here we conducted a reciprocal transplant experiment to examine how this variation in floral traits, under conditions of either high and low pollinator density, impacted seed production in the Tibetan lotus Saussurea nigrescens. In 2014 and 2015, we recorded the floral traits, pollinator visitation rates, and seed production of S. nigrescens populations grown in both home sites and foreign sites, where sites varied in honey bee population density. Our results demonstrated that the floral traits reflected those of their original population, regardless of their current location. However, seed production varied with both population origin and transplant site. Seed number was positively correlated with flower abundance in the pollinator‐rich sites, but with nectar production in the pollinator‐poor sites. Pollinator visitation rate was also positively correlated with flower number at pollinator‐rich sites, and with nectar volume at pollinator‐poor sites. Overall, the local genotype had higher seed production than nonlocal genotypes in home sites. However, when pollen is hand‐supplemented, plants from pollinator‐rich populations had higher seed production than plants from pollinator‐poor populations, regardless of whether they were transplanted to pollinator‐rich or ‐poor sites. These results suggest that the plant genotypic differences primarily drive variation in pollinator attraction, and this ultimately drives variation in seed: ovule ratio. Thus, our results suggest that flowering plant species use different reproductive strategies to respond to high or low pollinator densities.     

Effect of Seed Source, Topsoil Removal, and Plant Neighbor Removal on Restoring California Coastal Prairies

Grasslands are hot spots of biodiversity but are now widely threatened by changes in patterns of disturbances, such as grazing and fire regimes, exotic species invasions, and cultivation. The goal of this experiment was to find the most appropriate combination of treatments to reintroduce Danthonia californica, a formerly dominant perennial bunchgrass, into degraded California coastal prairies. Danthonia californica was sown from seed and transplanted at two sites and at two grazing intensities (grazed/ungrazed) in a multifactorial experiment testing the effects of (1) local versus nonlocal seed sources; (2) topsoil removal; and (3) reduction of plant neighbors. Seed emergence was very low, suggesting that transplanting may be a better option to reintroduce D. californica. Although transplants grown from nonlocal seeds survived better initially at both sites, transplants from local seeds had higher survival after 1.5 year at one site. This suggests that short‐term plant establishment studies may be misleading. Topsoil removal greatly enhanced transplant survival, and neighbor removal primarily increased transplant growth. Our results suggest that removing topsoil prior to transplanting seedlings grown from local seeds is the most promising method to reintroduce D. californica. However, the benefits of removing topsoil to provide safe sites for plant establishment should be weighed carefully against potential negative effects on the native seed bank and microbial communities on a site‐specific basis .     

Fitness in Naturally Occurring and Restored Populations of a Grassland Plant Lychnis flos‐cuculi in a Swiss Agricultural Landscape

Wildflower seed mixtures are widely used for restoration of grasslands. However, the genetic and fitness consequences of using seed mixes have not been fully evaluated. Here, we studied the role of genetic diversity, origin (commercial regional seed mixtures, natural populations), and environmental conditions for the fitness of a grassland species Lychnis flos‐cuculi. First, we examined the relationship between genetic diversity, environmental parameters, and fitness in sown and natural populations of this species in a Swiss agricultural landscape. Second, we established an experiment in the study area and in an experimental garden to study the implications of local adaptation for plant fitness. Third, to examine the response of plants to different soil properties, we conducted an experiment in climate chambers, where we grew plants from sown and natural populations of L. flos‐cuculi as well as from seed suppliers on soils with different nutrient and moisture content. We detected no significant effect of genetic diversity on the fitness of sown and natural populations. There was no clear indication that plants from natural populations were better adapted to local environment than plants from sown populations or seed suppliers. However, plants of natural origin invested more into generative reproduction than plants from sown populations or seed suppliers. Furthermore, in the climate chamber, plants originating from natural populations tended to flower earlier. Our results indicate that using nonlocal seeds for habitat recreation may influence restoration success even if the seeds originate from the same seed zone as the restored site.     

Effects of litter removal and mowing on germination and establishment of two fen-grassland species along a productivity gradient

To study the factors that limit the occurrence of species in fen grasslands, the effects of mowing and litter removal on germination, growth and establishment of two common species (Silene flos-cuculi andLotus pedunculatus) were analyzed along a productivity gradient at seven sites in Northern Germany. The sites differed in nutrient availability, vegetation composition and standing crop. In autumn 2002 a field experiment was set up at each site, including factorial combinations of two treatments (litter removal, mowing). Seeds and juvenile individuals of both species were transferred within the treatment combinations. The number of germinated and transplanted individuals was recorded during the 2003 and 2004 seasons and biomass of survived plants weighed in late summer 2004. Mowing and litter removal had positive effects on the number of germinated seeds of both species. The effect of litter removal was mostly significant at sites with a comparatively low nutrient availability and standing crop (350 g/m²), while the effect of mowing was more important at the site with the highest standing crop (1000 g/m²). In some cases the presence of litter had a positive effect on the survival of individuals, which may be caused by extremely dry weather conditions in 2003. The experiment showed that plant species reaction on mowing and litter removal differed both according to their life stage and the site conditions. These differences could be interpreted in context with Shifting Limitation Hypothesis because the abolition of seed limitation by sowing and creation of additional gaps for germination was most efficient at low productive sites, while the high standing crop reduced first of all the establishment probability, which could be increased by mowing.     

The spatial scale of adaptive population differentiation in a wide‐spread,well‐dispersed plant species

Adaptation to the specific conditions at different sites may contribute strongly to the wide distribution of a plant species. However, little is known about the scale at which such adaptation occurs in common species. We studied population differentiation, plasticity and local adaptation of the short‐lived perennial Hypochoeris radicata, a widespread and common plant whose seeds are well‐dispersed. We reciprocally transplanted seedlings among several populations of different size within and among three European regions (in the northwest Czech Republic, central Germany and the central Netherlands) and studied several fitness‐related traits over two growing seasons. The region in which the reciprocal transplant experiment was carried out had no influence on the performance of seedlings, indicating that there were no differences in overall habitat quality. In contrast, the site within region, and the plot within site strongly influenced mean plant performance. Plants from different populations of origin differed in their performance, indicating genetic variation among populations, but performance strongly depended on the specific combination of population of origin and transplant site. Plants that grew at their home site produced on average almost twice the number of seeds per transplant (a multiplicative fitness measure) than foreign plants originating from other sites. Survival, rosette size and multiplicative fitness all decreased with increasing distance from the home site to the transplant site. The size of the population of origin did not influence overall plant performance or the strength of local adaptation. In conclusion, our results indicate that the common and widespread H. radicata consists of locally adapted genotypes within its European range at a relatively small scale. Thus a large potential for gene flow by seeds and a high density of populations do not appear to be sufficient to prevent population differentiation by selection.     

Establishment of arbuscular mycorrhizal plants originating from xerothermic grasslands on heavy metal rich industrial wastes–new solution for waste revegetation

Industrial waste substrata, rich in heavy metals, are poorly suited for plant growth. Efforts are made to establish an appropriate plant cover to reduce erosion and further contamination. Grasses are the usual solution, as they grow fast, thrive on poor substrata and have well-developed root systems. Some of them are also highly dependent on mycorrhizal symbiosis that supports their growth especially on poor and polluted soils. However, the commercially available grasses often meet a lack of well established mycorrhiza on the site and the introduced plant populations dramatically decrease with time, despite large financial input including covering the substratum with soil and intensive watering. The aim of this paper was to select proper plants together with mycorrhizal fungi that could accelerate the establishment of the vegetation and improve its diversity under these extreme conditions, minimizing the financial costs of the reclamation (no use of soil layering and watering). The experiments were carried out under field and laboratory conditions. The plant seeds used originated from dry calcareous grasslands. The seeds were germinated under field conditions or in pots filled with soil supplemented with substratum from the industrial wastes. The seedlings were inoculated with AM fungi and introduced on the field plots a few weeks after germination. The inoculum consisted of either crude inoculum harvested from the dry calcareous grasslands or strains originating from polluted areas. Plants colonized by mycorrhizal fungi established well in the experimental plots. The results suggest that inocula from dry calcareous grasslands are potentially useful in revegetation of industrial wastes. Although in several cases the photosynthetic activity of plants was lower than at the natural sites, almost all plants survived and formed seeds. In all experiments the plant vitality was estimated on the basis of chlorophyll a fluorescence and was useful to show differences between waste substrata, inocula and coexisting plant species. The interactions between mycorrhizal and non-mycorrhizal plants were studied under greenhouse conditions and at least no negative effect of this coexistence was found.     

Experimental assessment of biotic and abiotic filters driving community composition

Species occurrence in a site can be limited by both the abiotic environment and biotic interactions. These two factors operate in concert, but their relative importance is often unclear. By experimentally introducing seeds or plants into competition‐free gaps or into the intact vegetation, we can disentangle the biotic and abiotic effects on plant establishment. We established a seed‐sowing/transplant experiment in three different meadows. Species were introduced, as seeds and pregrown transplants, into competition‐free gaps and the intact vegetation. They included 12 resident plants from the locality and 18 species typical for different habitats. Last two years, gaps were overgrown with vegetation from surrounding plants and we observed the competitive exclusion of our focal plants. We compared plant survival with the expected occurrence in target locality (Beals index). Many of the species with habitat preferences different from our localities were able to successfully establish from seeds and grow in the focal habitat if competition was removed. They included species typical for much drier conditions. These species were thus not limited by the abiotic conditions, but by competition. Pregrown transplants were less sensitive to competition, when compared to seedlings germinated from seeds. Beals index significantly predicted both species success in gaps and the ability to withstand competition. Survival in a community is dependent on the adaptation to both the abiotic environment and biotic interactions. Statistically significant correlation coefficients of the ratio of seedling survival in vegetation and gaps with Beals index suggest the importance of biotic interactions as a determinant of plant community composition. To disentangle the importance of abiotic and biotic effect on plant establishment, it is important to distinguish between species pool as a set of species typically found in given community type (determined by Beals index) and a set of species for which the abiotic conditions are suitable.     

Restoration of ditch bank plant species richness: The potential of the soil seed bank

Abstract. Small‐scale landscape elements, such as ditch banks, play an important role in preserving plant species richness in agricultural landscapes. In this study, we investigate whether the seed bank might be useful for restoring the above‐ground plant species richness. We studied the vegetation and seed bank composition at six species‐rich and six species‐poor ditch banks, where agri‐environment schemes are running to maintain and enhance ditch bank plant diversity. We show that the number of species in the seed bank was low, regardless of the number of species in the established vegetation. Moreover, the seed bank was always dissimilar to the established vegetation. Target species for nature conservation were occasionally present in the seed bank at both species‐poor and species‐rich sites, but rarely so if the species was absent from the established vegetation. We conclude that the potential use of the seed bank for restoration of ditch banks is minimal. At present, plant species richness seems to be largely controlled by germination opportunities; high biomass and competition appear to hamper germination at species‐poor sites. We recommend continued nutrient reduction at such sites. Soil disturbance measures and deliberate sowing should also be considered.     

Genetic Diversity and the Reintroduction of Meadow Species

Abstract: Restoration of formerly nutrient‐poor and species‐rich grasslands generally leads to an increase in species diversity. However, species without a persistent seed bank and with poor dispersal ability often do not re‐establish spontaneously. Here, reintroduction is an option. If existing populations are comparable in their genetic composition, any population will do. This is not the case if populations have local adaptations. Unfortunately, whether populations are adapted locally is not easily determined, in contrast to assessing differentiation using neutral genetic markers. We used AFLP to study genetic diversity of Cirsium dissectum and Succisa pratensis within and among several Junco‐Molinion plant communities in the Netherlands (up to 200 km apart) that were potential source populations, and followed the reintroduction using seeds from these populations. Also, vegetative growth phase characteristics of three populations of C. dissectum were analyzed under controlled conditions. Most of the genetic variation in these cross‐fertilizing species was found within populations. Small but significant genetic differences in band frequencies were found among populations (F_st 0.100 ‐ 0.135). The first generation of reintroduced plants contained less polymorphic bands than the source populations. The genetic differences caused by reintroduction using a limited number of seeds (founder effects) were significant in all except one case (F_st 0.012 ‐ 0.101 between source and corresponding reintroduced population), but the magnitude was smaller than the source population differentiation. In assignment tests, reintroduced populations resembled their source population more than any other population, but all populations contained sizeable proportions of plants that were assigned to most similar plants from other populations, indicating that the populations are only marginally distinct. Calculations show that reintroduction from more than one source population introduces significantly more polymorphic bands into the new population, capitalizing on the existence of band frequency differences among populations.     

Effects of Habitat Quality and Isolation on the Colonization of Restored Heathlands by Butterflies

In areas with intensive land use, such as the Netherlands, habitat fragmentation and loss of habitat quality due to eutrophication and drainage are major threats to the preservation of species‐rich communities of heathland and acid grassland. Restoration of such nutrient‐poor habitats may be carried out by removing the topsoil from ex‐arable land, in order to lower the nutrient levels. However, the establishment of target plant communities is known to be fragmentary. The current study shows that this also applies to butterflies. Ten years after topsoil removal in eight study areas, on average, only 3.5 of 10 characteristic heathland species were recorded on the sites. Species that did colonize had a significantly lower density than in the source populations. Our study indicates that although isolation effects were limiting colonization, poor habitat quality was the main limiting factor, mainly due to lack of host plants, hydrological conditions, and, to a lesser extent, lack of nectar plants and excessive residual nutrient levels. An experiment with the introduction of cut heather in one study area showed a significantly higher abundance of both target and nontarget butterflies in manipulated sites than in control sites. It can be concluded that habitat restoration by topsoil removal can be successful for butterflies of especially wet heathland habitats, provided that source populations are at close range and care is taken that complete plant communities are restored.     

Investigating Predictors of Plant Establishment During Roadside Restoration

Ecological restoration on roadsides confers several ecological benefits, but also poses significant challenges. Native plants used in restoration efforts must survive compacted soil, harsh microclimates, prolific invasive species, and pollution from road salts and vehicle emissions. Criteria for both site and species selection need to be developed to assist practitioners in restoring roadside environments. We transplanted seedlings of 9 grassland plant species into plots within 8 highway interchanges surrounding Ann Arbor, Michigan, United States. To see if they might predict plant performance in roadside restoration, we assessed 2 indices: the coefficient of conservatism (CC) as an index of affinity to undisturbed habitat; and the number of U.S. counties in which each species occurs as an index of distribution. We measured seedling survival, height, and biomass during the first growing season, and survival 1 year after transplant. We also measured soil characteristics, air temperature, and humidity at each interchange. We found that soil characteristics largely determined plant survival. Plants were more likely to survive in sandier soils than in soils rich in silt or clay that had high bulk density, high pH, and high conductivity. Although plant survival varied among species and interchanges, neither CC nor county‐level distribution was a useful predictor of survival. Our results illustrate the importance of matching plant species with local soil characteristics when choosing restoration sites, and offer guidance to transportation officials considering roadside restoration with native plants, and practitioners working to restore any heavily disturbed site.     

Restoration Ecology of an Endangered Plant Species: Establishment of New Populations of Cirsium pitcheri

We determined the effects of shade, burial by sand, simulated herbivory, and fertilizers on the survival and growth of artificially planted population of Cirsium pitcheri—an endangered plant species of the sand dunes along Lake Huron. Sand burial experiments showed that greenhouse grown plants should optimally be transplanted into areas receiving 5 cm of sand deposition: burial at this depth maximized emergence, survivorship, and below‐ground biomass. Under field conditions, simulated herbivory of up to 50% of the plant height produced a slight increase in biomass after one year of growth. Field observations showed that when white‐tailed deer removed more than 50% of the transplant's leaf tissue, the plant died. The application of a 20:20:20 (N:P:K) water‐soluble fertilizer produced a significant increase in the dry leaf biomass, total leaf area, and total dry biomass relative to control plants. We also tested for the presence or absence of a persistent seed bank. Few seeds were recovered from soil samples collected from Pinery Provincial Park and Providence Bay. However, C. pitcheri has the ability to form a persistent seed bank under field conditions but only at soil depths of 15 cm. Cirsium pitcheri seeds are able to germinate and seedlings can emerge from a burial depth of up to 6 cm. Thus, seeds planted in open, sunny areas will probably maximize emergence, growth, and survivorship of seedlings. Populations of C. pitcheri can be restored by planting seeds at shallow depths, transplanting greenhouse‐grown plants, applying water soluble fertilizers, and protecting plants from herbivores.     

Restoration of Species‐Rich,Nutrient‐Limited Mountain Grassland by Mowing and Fertilization

Mowing and management to reduce nutrient levels have often been successfully used to restore species‐rich grasslands in various parts of Europe. However, such treatments have failed to restore the species‐rich Central European mountain grasslands dominated by Polygonum bistorta. P. bistorta builds an extensive underground rhizome system that monopolizes available nutrients in these nutrient‐poor grasslands, enabling this species to persist at high densities even in the presence of mowing. Therefore, we tested a restoration approach using a factorial combination of fertilization and mowing, as well as a litter removal treatment. The experiment was run over 5 years and species composition response to these treatments was recorded at two spatial scales. Mowing suppressed flowering and cover of P. bistorta and promoted target grassland species and richness. Fertilization prevented nutrient impoverishment and increased height and dominance of the broad‐leaved grasses with which many species‐rich grassland herbs could coexist. The additive effect of the mowing/fertilization treatments was strong enough to act as a driver of P. bistorta suppression and associated community change. The litter removal treatment, however, had little effect on plant composition. The experiment demonstrates that in nutrient‐limited grasslands, increasing nutrient levels in addition to mowing to manage competition for light can be used to control dominants. This contrasts with restoration of systems where after abandonment increased nutrient levels lead to the establishment of tall dominants that suppress other species by competition for light.     

Sowing and transplantation of Adonis ramosa (Ranunculaceae) for the establishment of a new population in a seminatural broad-leaved deciduous forest floor

Adonis ramosa (Ranunculaceae) is a plant whose risk of extinction is increasing in Japan. In this study, we show that there is a significant possibility of establishing an A. ramosa population by sowing seeds or transplanting plants in a seminatural broad-leaved deciduous forest floor. Immediately after seed collection in early June, seeds were sown either in pots that were watered periodically or in a deciduous forest floor where the soil surface was prepared. In the following spring, 30–40% of the seeds germinated. Of the one-year-old plants buried at a depth of 2 cm and two-year-old plants buried at a depth of 2 or 8 cm in the fall, 96% sprouted leaves in the following spring. However, 79% of the one-year-old plants buried at a depth of 8 cm sprouted leaves. In spring, the number of leaves, the widths of individual plants, and the heights of the two-year-old plants were greater than those of the one-year-old plants. The dry weights of one- and two-year-old plants in fall after one spring growing season increased by 1.6–2.3 times and by 2.3–5.1 times, respectively, compared to the dry weight at the time of transplanting. Also, the dry weights of the plants transplanted at a depth of 8 cm were higher than those of plants transplanted at a depth of 2 cm for both one- and two-year-old plants. The results suggest that the growth conditions at a depth of 8 cm are more favorable than those at a depth of 2 cm.     

Restoring grasslands with arbuscular mycorrhizal fungi around remnant patches

Ecological restoration of species‐rich grasslands remains a priority for conservation of biodiversity. Torrez et al. (Applied Vegetation Science, this issue) determined if plant species recolonization of degraded nutrient‐poor grasslands could be increased by adding a local source of arbuscular mycorrhizal fungal (AMF) inoculum at different distances from intact remnant grasslands. They highlight the important role of below‐ground processes on restoration success.     

Late Transplanting and the Yield of Phaseolus vulgaris L.

The prediction that very high seed yields of dry beans (Phaseolusvulgaris L.) would be produced by the delayed transplantingof large plants has been tested in a factorial experiment withfour dates of transplanting and eight plant populations. Therewere significant differences in yield between transplantingdates and between population densities, and there was a significantdate-density interaction. At low plant densities (up to about30 plants m^–2) the three transplanted treatments yieldedless than the hand-sown controls, and late transplanting yieldedless than early. At the highest density the situation was reversed;all three transplanted treatments out-yielded the controls andlate transplanting tended to out-yield plants transplanted early.The biggest yield was 340 g seed m^–2 from a transplantedcrop grown at 35 plants m^–2. The data on yield fitted a modified rectangular hyperbola ofthe form where y is yield per unit area, p is the number of plants perunit area, t is the number of days between sowing and transplanting,and B_o, n, m, and p are arbitrary parameters. This equationaccounted for 91 per cent of the variation in yield with t andp. It is suggested that late transplanting had adverse effects,due to transplanting ‘shock’ and which were mostmarked at low plant densities; and beneficial effects, ascribableto an effect on plant ‘plasticity’, which were mostmarked at high plant densities. Possible physiological mechanismsof these effects are discussed. Phaseolus vulgaris, yield, density, transplanting     

Reestablishment of the Southern California Rocky Intertidal Brown Alga,Silvetia compressa: An Experimental Investigation of Techniques and Abiotic and Biotic Factors That Affect Restoration Success

Previous research has indicated that many rocky intertidal macrophyte communities in southern California, and other locations around the world, have shifted from larger, highly productive, fleshy seaweeds toward a smaller, less productive, disturbance‐tolerant flora. In widespread decline are ecologically important, canopy‐forming, brown seaweeds, such as the southern California rockweed species Silvetia compressa. Restoration efforts are common for depleted biogenic species in other habitats, but restoration within rocky intertidal zones, particularly on wave‐exposed coasts, has been largely unexplored. In two phases, we attempted to restore Silvetia populations on a southern California shore by transplanting live plants and experimentally investigating factors that affect their survival. In Phase I, we implemented a three‐way factorial design where juvenile Silvetia thalli were transplanted at four sites with a combination of simulated canopy and herbivore exclusion treatments. Transplant survival was low, although enhanced by the presence of a canopy; site and herbivore presence did not affect survival. In Phase II, we used a two‐way factorial design, transplanting two size classes of rockweeds (juveniles and reproductive adults) on horizontal and partially shaded, north‐facing vertical surfaces at a target location where this rockweed has been missing since at least the 1970s. Transplant survival was moderate but lower than natural survival rates. Larger thalli exhibited significantly higher survival rates than smaller thalli in both the transplanted and naturally occurring populations, particularly on vertical surfaces. Higher mortality on horizontal surfaces may have been due to differences in desiccation stress and human trampling. Transplanting reproductive adults resulted in the subsequent recruitment of new individuals.     

The influence of seed addition and cutting regime on the success of grassland restoration on former arable land

Questions: Can seed addition enhance the success of establishing species‐rich grassland on former arable land? Are sowing date and cutting regime important in determining success? Location: Aberdeen and Elgin, northeast Scotland, United Kingdom. Methods: A field experiment was conducted at two sites to assess the effect of seed addition, sowing date and cutting regime on the vegetation developing on former arable land, the aim being to compare the success of different treatments at producing a species‐rich grassland. Results: Sowing a seed mix resulted in the establishment of vegetation very distinct from the species‐poor vegetation dominated by perennial grasses which otherwise developed, though establishment success of the sown grassland species was highly variable between sites. Where establishment of the sown species was poor, sowing date had no significant effect on species composition, whereas the cutting regime was very important. Cutting the vegetation significantly increased both the number and abundance of sown species compared with the uncut control. Conversely, where establishment had been good, the cutting regime in the first year had little effect on species composition. Cutting the vegetation at least twice a year appeared to be the most effective management over the length of the experiment. Conclusions: Sowing a seed mixture significantly reduced the abundance and number of naturally colonising species, effectively controlling problem weed species such as Senecio jacobaea and Cirsium vulgare, highlighting the agronomic value of sowing seed mixtures on fallow farmland. The sowing of a seed mix on former arable land has demonstrated that it is feasible to create vegetation similar in character to that of species‐rich grasslands.     

Radial Dispersion of Neighbors and the Small-Scale Competitive Impact of Two Annual Grasses on a Native Perennial Grass

In California's Mediterranean type grasslands, native perennial grasses such as Nassella pulchra are surrounded by introduced annual species and these annuals are thought to have displaced natives through much of their range. Amongst other invaders, two grasses Lolium multiflorum and Bromus hordeaceus, commonly dominate portions of the grassland with potential for N. pulchra restoration. We hypothesized that competitor species differences and small‐scale gaps (150 cm²) could be important determinants of N. pulchra survival and performance on these sites. Lolium multiflorum and B. hordeaceus were planted in 20 cm diameter circular plots at a constant rate of 1 seed per cm² surrounding newly transplanted N. pulchra plants. Nassella pulchra showed no significant effect of the species of competitor or from the distribution of the competitors. Both interspersion of patches of bare ground and separation of competitors into patches did not increase N. pulchra pre‐dawn water potential, basal area change, number of seeds produced, or average weight of seeds. The presence of L. multiflorum was associated with a decrease in N. pulchra survival compared with plots with only B. hordeaceus. Plants with increases in basal area of less than 0.75 cm² during the growing season had 74% mortality compared with no mortality in plants with more growth. However, initial N. pulchra plant size was not a good predictor of mortality. Limiting competition from annuals may increase survival of N. pulchra plantings, but 60% of the plants survived for at least 1 year, despite being transplanted into soil containing substantial annual grass seed.     

Mass effects,clonality, and phenology but not seed traits predict species success in colonizing restored grasslands

Preserved grasslands commonly host species‐rich plant communities but a large part of the grasslands were plowed up in the past. Their restoration often requires a long time and initial restoration measures might trigger ecosystem recovery, which is then followed by spontaneous colonization. We evaluate the establishment success of target grassland species, which were not sown but established spontaneously in the restored grasslands of Bílé Karpaty Mts., SE Czech Republic. According to their key functional traits and incidence in the landscape (mass effect; acquired from the results of a grid mapping project in the region), we examined the frequency of species and their mean cover in 82 restored grasslands. The best predictor of species frequency in the grasslands was their mass effect, followed by a high capacity for clonal growth and late phenology. Seed dispersal traits (seed mass, terminal velocity, epizoochory ranking index) and plant height had no significant effect. Specific leaf area was positively correlated with mass effect. Species having a high cover in the restored grasslands had a high capacity for clonal growth. In the preparation of seed mixtures, we should therefore consider that nonclonal species relying on regeneration from seeds will be generally less able to reproduce and should be promoted by artificial sowing. At the same time, species common in the landscape, which spread well clonally, and those with a late phenology, might be expected to colonize restored meadows on their own, so that sowing them is not necessary.