PREMISE OF THE STUDY: Vegetative reproduction and spread through clonal growth plays an important role in arctic-alpine ecosystems with short cool growing seasons. Local variation in winter snow accumulation leads to discrete habitat types that may provide divergent conditions for sexual and vegetative reproduction. Therefore, we studied variation in clonal structure of a dominant, evergreen, dwarf shrub (Empetrum nigrum s.l. with the two taxa E. nigrum L. s.s. and E. hermaphroditum Hagerup) along a snow cover gradient and compared clonal diversity and spatial genetic structure between habitats.

METHODS: We studied 374 individual shoots using 105 polymorphic AFLP markers and analyses based on hierarchical clustering, clonal diversity indices, and small-scale spatial genetic structure with pairwise kinship coefficient. We used two approaches to define a threshold of genotypic distance between two samples that are considered the same clone. Clonality was examined among three habitats (exposed ridges, sheltered depressions, birch forest) differing in snow conditions replicated in four study regions in Norway and Sweden.

KEY RESULTS: Clonality of E. hermaphroditum differed between habitats with an increase in clonal diversity with decreasing snow depth. Small-scale spatial genetic structure increased with decreasing clonal diversity and increasing clone size. In three study regions, E. hermaphroditum was the only species, whereas in one region E. nigrum also occurred, largely confined to exposed ridges.

CONCLUSIONS: Our results demonstrated that snow cover in conjunction with associated habitat conditions plays an important role for the mode of propagation of the dwarf shrub E. hermaphroditum.

Arctic ecosystems face strong changes in snow conditions due to global warming. In contrast to habitat specialists, species occupying a wide range of microhabitats under different snow conditions may better cope with such changes. We studied how growth and reproduction of the dominant dwarf shrub Empetrum hermaphroditum varied among three habitat types differing in winter snow depth and summer irradiation, and whether the observed patterns were consistent along a local climatic gradient (sub-continental vs. sub-oceanic climates) and along a latitudinal gradient (northern Sweden vs. central Norway). Habitat type explained most of the variation in growth and reproduction. Shoots from shallow snow cover and high summer irradiation habitats had higher numbers of flowers and fruits, lower ramet heights, shorter shoot segments, lower numbers of lateral shoots and total biomass but higher leaf density and higher relative leaf allocation than shoots from habitats with higher snow depth and lower summer irradiation. In addition, biomass, leaf allocation and leaf life expectancy were strongly affected by latitude, whereas local climate had strong effects on seed number and seed mass. Empetrum showed high phenotypic trait variation, with a consistent match between local habitat conditions and its growth and reproduction. Although study areas varied strongly with respect to latitude and local climatic conditions, response patterns of growth and reproduction to habitats with different environmental conditions were consistent. Large elasticity of traits suggests that Empetrum may have the potential to cope with changing snow conditions expected in the course of climate change.

The topography within arctic-alpine landscapes is very heterogeneous, resulting in diverse snow distribution patterns, with different snowmelt timing in spring. This may influence the phenological development of arctic and alpine plant species and asynchronous flowering may promote adaptation of plants to their local environments.We studied how flowering phenology of the dominant dwarf shrub Empetrum hermaphroditum varied among three habitats (exposed ridges, sheltered depressions and birch forest) differing in winter snow depth and thus snowmelt timing in spring, and whether the observed patterns were consistent across three different study areas.Despite significant differences in snowmelt timing between habitats, full flowering of E. hermaphroditum was nearly synchronous between the habitats, and implies a high flowering overlap. Our data show that exposed ridges, which had a long lag phase between snowmelt and flowering, experienced different temperature and light conditions than the two late melting habitats between snowmelt and flowering.Our study demonstrates that small scale variation seems matter less to flowering of Empetrum than interannual differences in snowmelt timing.

Regularly mown road verges are an important habitat for conservation of grassland vegetation. Disturbance and movement of seed-contaminated soil during road construction and maintenance makes road verges susceptible to the establishment of invasive alien plants such as garden lupine (Lupinus polyphyllus Lindl.). To combat spread of L. polyphyllus via seeds, we tested methods for seed destruction using heat. This study aimed at developing heat eradication methods for dry and imbibed L. polyphyllus seeds applying dry heat (88, 93, 98, 103 C at 1, 3, 5, 10 min) in a laboratory, steam (85, 90, 95 C at 3, 5, 10 min) in a test-box steaming device and (97 C at 10-17 min; dry seeds only) in a stationary soil-steaming machine (S30). In order to speed up water absorption and post-treatment germination, the imbibed seeds were manually scarified before the heat treatment and the dry seeds afterwards. Additionally, germination of two different age seed batches was tested applying dry heat (88, 98 C at 3, 5 min). The results showed that steam treatments inhibited seed germination more than dry heat in both dry and imbibed seeds. Germination dropped to < 5% when steamed at ≥ 90 C or dry-heated at > 100 C. Seed germination decreased with higher temperatures and longer exposure times. Imbibed seeds exhibited lower germination compared to dry seeds for dry and steam heat. Approximately 0.5% of dry seeds germinated when steamed using S30. 2022-collected seeds were less sensitive to dry heat th an seeds from 2020. In conclusion, hot steam is more effective in reducing L. polyphyllus seed germination than dry heat. Thus, to successfully eradicate L. polyphyllus seeds in soil masses, we recommend steaming them at 97 C for at least 10 minutes. 

The dynamics of many plant populations essentially depend upon seed and seedling stages, and a persistent seed bank may give species an opportunity to disperse through time. Seed burial is a decisive prelude to persistence and may strongly influence seed-bank dynamics. The fate of buried seeds depends on species-specific traits, environmental conditions and possibly also burial mode. We tested seed germination, seedling emergence and growth of the co-occurring herbaceous flood-meadow species Arabis nemorensis, Galium wirtgenii, Inula salicina, Sanguisorba officinalis and Selinum carvifolia in response to the experimental manipulation of burial depth (0, 1, 2, 4, 8, 12 cm) and substrate type (sand, clay). Increasing burial depth led to decreased germination, emergence and growth in all species studied, and seedling growth differed significantly between substrate types. The responses of species differed on an individual basis, but also showed a higher-ranking pattern based on seed size. Larger-seeded species were able to emerge from greater depths and experienced less depth-mediated growth inhibition than smaller-seeded species, which, in turn, had higher survival rates during burial and were less likely to experience fatal germination. Based on these results, we suggest that herbaceous flood-meadow species have developed two different seed-size based strategies for coping with the extreme recruitment conditions prevailing in flood meadows, the balance of which seems to be maintained by disturbance events.

The main objective of this study was to analyze whether plant material transfer is a suitable approach for establishing plant populations with spatial patterns and population structures resembling those of remnant populations. We studied pattern formation and population characteristics in three remnant and two restored populations of the biennial plant species Arabis nemorensis in the upper Rhine valley in southwestern Germany over a period of 2 years. We investigated spatial patterns of seedlings, juveniles, and adults as well as the small-scale horizontal and vertical distribution of seeds in the soil, followed the fate of individual plants and recorded structural habitat parameters such as vegetation and litter cover. Population dynamics differed between the study sites, but there was no pronounced difference between restored and remnant sites. Seedlings, juveniles, and adults as well as seeds in the seed bank showed aggregated spatial patterns on all study sites, with positive autocorrelation on a scale of 20-60 cm. Within sites, patterns remained approximately stable through time. Restored sites experienced rapid seed bank formation as a result of the restoration measures. Our results suggest that the restoration measures were not only successful in transferring the target species but also triggered rapid formation of spatially structured populations that, years after restoration, closely resembled those of remnant sites.

Desiccation cracks are a natural phenomenon of clay-rich soils that form via soil shrinkage during dry conditions. Our aim was to test the seed trapping potential of such cracks and assess its impact on seed bank formation in a flood-meadow ecosystem. We documented crack patterns on permanent plots and analysed the soil seed content along and adjacent to cracks. Seed translocation via cracks was tested with a mark-recapture experiment, and post-entrapment seed fate was tested with a burial experiment. Most cracks re-opened in the same positions in consecutive dry periods. Along cracks, most seeds were found in 10-20 cm depth, whereas adjacent to cracks most seeds were found in 0-5 cm depth. The majority of seeds found in shallow depths adjacent to cracks belonged to species that were also present in the above-ground vegetation, whereas this rate was always under 50% along desiccation cracks. The mark-recapture experiment gave evidence for vertical seed translocation through desiccation cracks. Post-entrapment seed fate differed between species and burial depth, with a trend towards increasing survival with increasing depth. We conclude that desiccation cracks act as natural seed traps, foster seed bank formation and thus influence plant community dynamics in flood meadow systems.

Plant material transfer is a well-established technique for overcoming dispersal limitation during grassland restoration. As restoration sites are frequently more abundant than donor sites, the plant material is often applied as patches or strips, with the assumption that these will act as colonization initials from which transferred species will spread and eventually cover the entire sites. Our aim was to test this assumption and to evaluate whether it is feasible to restore entire sites by spatially-restricted plant material application in a flood-meadow ecosystem. We established transverse transects consisting of eight 2 x 2 m plots on five plant material strips 7-8 years after plant material application. We monitored the above-ground vegetation development, analyzed the seed rain and determined the composition of the soil seed bank, i.e. we compared three different components of the emerging flood-meadow community. Transferred species were present in all three community components studied, and 88.6% of the 79 species we found in total had already spread from the plant material strips and colonized their surroundings. Detected dispersal distances differed between community components, and the share of colonizers was highest for the above-ground vegetation and lowest for the soil seed bank. We conclude that plant material transfer is a suitable technique for restoring flood-meadows as transferred species not only establish on the sites supplied with plant material, but also colonize their surroundings. (C) 2010 Elsevier Ltd. All rights reserved.

Most range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites. We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded. We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance. Our results suggest that abiotic and biotic soil characteristics can shape climate change-driven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position x earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.

Establishment of plants through seeds is often constrained by the quality of microsites, which is in part controlled by the nature and amount of ground cover. The latter consists of living shoots of vascular plants or bryophytes and/or the dead remains of the dominant species. In the present article, we report the results of a controlled pot experiment with five species characteristic of floodplain grasslands. We manipulated the amounts of grass litter and/or mosses to study (1) differences between ground cover types with respect to their effects on microenvironment and seedling emergence and (2) how these effects interact with seed size and seed sowing position. Increasing amounts of both cover types led to increasing soil humidity, whereas temperature amplitude and illumination were decreased. However, since grass litter decomposed much faster than bryophytes, light conditions for germination under grass litter improved considerably with time. Although seedling emergence varied significantly between species, ground cover types and cover amounts, seed position alone explained about 50% of the variation in the data set. Additionally, we found an important interaction between seed size, seed position and cover type: large-seeded species showed a fitness advantage when seeds were situated beneath a cover, irrespective of cover type, which disappeared when seeds were shed on top of a cover layer. We suggest that this interaction may be ecologically and evolutionarily relevant because it may lead to changes in species composition and diversity of plant communities as a consequence of changes in the amount and type of ground cover.

1. The effects of litter on seedling emergence (e.g. during succession from grassland to woodland) may not only depend on litter amount and environmental conditions but may also be related to litter types. We tested the effect of litter types, litter cover and soil moisture on the emergence of four congeneric pairs of grassland and woodland species. 2. We carried out a controlled pot experiment (Experiment I) using litter and species from two habitats (grassland vs. woodland), three levels of litter cover (low, medium, high) and two levels of water-addition (leading to constantly humid or intermittently dry substrate). Amounts of litter were adjusted to result in the same relative light reduction for both litter types. Consequently, we applied 2 g, 4 g and 8 g of grassland litter and 0.5 g, 1 g and 2 g of woodland (oak) litter per pot. 3. To separate the shade effect of litter from its mechanical plus chemical effects we exposed additional pots with seeds of the same species to deep shade conditions (Experiment II) corresponding to 4 g and 1 g of grass and oak litter, respectively. 4. Under intermittently dry conditions both litter types increased seedling emergence. In addition, we found a significant interaction between litter type and species origin: seedling emergence of woodland species was significantly lower from beneath grass litter than from beneath oak litter, whereas grassland species emerged equally well from beneath both litter types. Compared with seed germination under a shade cloth, seedling emergence of woodland species from beneath grass litter was reduced by 44%. Litter significantly affected soil humidity and the amplitude of diurnal temperature fluctuations. Differential effects of litter types on woodland and grassland species are probably related to litter structure. 5. Synthesis. Our experimental data present evidence that ecosystem specific litter effects slow down succession from grassland to woodland and that the effect size is controlled by the litter amount present and the environmental conditions.

Fitness of individual plants and of populations depends on the rates of survival, growth and fecundity. This study tested whether vital rates were differentially affected by biotic interactions and water availability. The effects of manipulations of above-ground competition (through clipping) and water availability (through water addition) on the vital rates of seedlings of three species (Viola elatior Fries, Viola pumila Chaix and Viola stagnina Kit.) were analysed in dry, mesic and wet grasslands. Water addition and grassland type had the largest effects on survival (accounting for 41 and 24% of total variation, respectively) across species. Height growth rate was positively affected by grassland type (19%) and water addition (12%) and varied among species (8%), while leaf accumulation rates and reproduction were affected by grassland type and clipping. The data suggested facilitative effects of the canopy on seedling survival in the dry grassland. This study presents evidence that environmental conditions and biotic interactions may have differential effects on seedling survival, growth and reproduction. The findings highlight the complex interplay between spatial and temporal environmental variation and biotic interactions in structuring plant communities.

We studied the demography of Viola elatior, V. pumila, and V. stagnina, three rare and endangered Central European floodplain species, to (i) analyse variation in life-cycles among congeners and between regions (Dyje-Morava floodplains, Czech Republic; Upper Rhine, Germany), (ii) to define sensitive stages in the life-cycles, and (iii) to identify possible threats for population viability and species conservation. Matrix models were based on the fate of marked individuals from a total of 27 populations over two years. We analysed population growth rate (lambda), stage distribution, net reproductive rate (R (0)), generation time, age at first reproduction, and elasticity and calculated a life table response experiment (LTRE). Most populations were declining and lambda did not differ between species or regions during the observed interval. Despite higher probabilities for survival and flowering in the Dyje populations, R (0) was higher in the Rhine populations. Also other demographic traits showed consistent differences between regions and/or species. Complex life-cycles and large variation in lambda precluded unequivocal identification of sensitive stages or vital rates for conservation. Variation between regions may be a consequence of differences in habitat quality. Our results suggest that deterministic processes such as reduced management, succession, habitat destruction, and lack of disturbance through reduced or eliminated flooding present the strongest threat for the viability and persistence of populations of the three floodplain violets as compared with stochastic processes. However, the persistent seed bank of the species may buffer populations against environmental variation and represents a reservoir for recovery after resumption of suitable land-use management.

We studied seedling emergence in four familial pairs of floodplain herbs in response to the experimental manipulation of soil moisture and litter cover to analyse (i) whether the effect of litter changes from negative under humid to positive under dry conditions, and (ii) whether the response to changing water and light conditions with increasing litter cover varies among species and plant families. We carried out a controlled pot experiment using four levels of litter cover (0 g, 2 g, 4 g and 8 g litter per pot, corresponding to 0 kg m(-2), 0.2 kg m(-2), 0.4 kg m(-2) and 0.8 kg m(-2)) and two levels of water-addition, leading to constantly humid substrate or intermittently dry topsoil. Across water-additions, percentage emergence reached a peak at low levels of litter cover (0.2 kg m(-2) and 0.4 kg m(-2)). There was a significant litter x water-addition interaction in six species, with positive effects of litter under intermittently dry conditions and negative or neutral effects under constantly humid conditions. Litter lowered maximum temperature as well as amplitude, and alleviated soil humidity under low water supply, while imposing increasingly shaded conditions. Analysis of species- and family-specific responses suggested that germination under a litter cover of 0.8 kg m(-2) was significantly reduced in smaller-seeded species (i.e. those that tend to have higher light demands for germination). Our results suggest that transfer of seed-containing plant litter can aid restoration projects if applied at 0.2-0.4 kg m(-2). Below these levels, establishment of most species may be inhibited by drought, while higher amounts will increasingly suppress seedling emergence, especially of small-seeded species. In addition to facilitation effects observed between living plants, dead plant remains may also exert positive effects on establishment. The sign of the litter effect on seedling emergence depends on soil humidity, with negative effects seen above a threshold amount, which is species- and family-specific and is closely related to seed size. Whether positive litter effects in grasslands are a consequence of coevolution remains to be examined.

Viola elatior, V pumila and V stagnina (syn. V persicifolia) are very rare in central and western Europe and red-listed in many central European countries. To improve their conservation and to base management efforts on sound scientific knowledge, we here review the available information on their biology. A comparative approach is adopted to identify similarities and differences between the species. Special emphasis will be on taxonomy, community ecology and population biology. All three species belong to the section Viola, subsection Rostratae. Since they are morphologically rather similar and have often been misidentified, we supply the most important characters for separating the three taxa. The species share a continental distribution with a centre of occurrence in the temperate zone of eastern Europe and western Siberia, and reach their western range margin in central and western Europe. They have become rare and endangered through melioration and fragmentation of their habitats. All three species are iteroparous hemicryptophytes with a complex life cycle, a mixed mating system with chasmogamous and cleistogamous flowers, and a persistent seed bank. Viola pumila and V. stagnina occur in floodplain meadows and wet grasslands, whereas V. elatior is a typical species of alluvial woodland fringes and other ecotonal habitats bordering floodplain meadows. Viola elatior and V pumila are confined to calcareous or at least base-rich substrates, whereas V stagnina may also occur on strongly acidic soils. The violets are able to occupy a broad range of site conditions in terms of soil nutrient status and productivity. However, on fertile sites the species depend on regular disturbances that weaken competitors and enhance the germination of dormant seeds. Current information about mycorrhizal colonisation, biochemical and physiological data, and herbivores and pathogens is scarce or lacking, probably due to the rarity of the three species in central Europe. (c) 2006 Rubel Foundation, ETH Zurich. Published by Elsevier GmbH. All rights reserved.

Physiological integration in clonal plants, which can be assumed to be dependent on vascular connections among ramets, is associated with several potential benefits, especially in nutrient-poor environments. However, some experimental evidence indicates that ectohydric bryophytes, i.e. species lacking specialised tissues for internal water conduction, also have physiologically integrated ramets. We tested this hypothesis by analysing nitrogen dynamics and tracing movements of a ¹⁵ N label among interconnected ramets of the ectohydric Hylocomium splendens over one season. The observed patterns were compared with translocation patterns in Polytrichum commune, an endohydric species that is known to show a high degree of clonal integration. Our aims were (1) to evaluate the degree of physiological integration among segments in H. splendens and (2) to study whether the pattern of ¹⁵ N movement obtained matched those depicted by changes in total nitrogen pool size. Current-year segments (G0) of both species were identified as strong sinks for nitrogen owing to their considerable increase in the ¹⁵ N pool during the season. In P. commune all other segments types showed a net loss of ¹⁵ N from June to September, which was probably due to autumn resorption of nitrogen to subterranean structures. In H. splendens one-year-old segments (G1) increased their ¹⁵ N pool, while older green segments (G2+) lost 50% of their initially absorbed ¹⁵ N. All the label lost from these source segments could be recovered in G0 and G1 segments. We suppose that most of the recycled nitrogen is provided by degeneration of three-year-old segments, which turn brown in parallel with the reallocation of nitrogen during the season. The high degree of physiological integration in H. splendens is discussed with respect to its life history and ecosystem nitrogen cycling.

Life history theory predicts that reproduction incurs costs in terms of future growth and survival. In infertile environments reproductive events may accelerate nutrient turnover such that losses of limiting nutrients cannot be replenished by uptake from the soil. 

We compared the effect of reproduction on nitrogen (N) turnover and N use-efficiency, i.e. the annual dry matter produced per unit N lost, of three carnivorous species of the genus Pinguicula on an infertile subarctic heath. We hypothesized that reproduction should increase N turnover, unless a larger N pool or improved N resorption can compensate for the losses associated with reproduction.

These carnivorous herbs showed a nitrogen use strategy similar to that of non-carnivorous plants (herbs and grasses) with a relatively low mean residence time (MRT) of N and a large dry matter productivity per unit N in the plant (annual N productivity). N pool size and the efficiency of N resorption from senescent leaves were similar in reproductive and non-reproductive individuals. Reproductive individuals had significantly larger annual N losses and thus a lower average MRT (0.8 vs. 1.8 years), with values < 1 indicating that the annual N losses are greater than the annual average N pool. Unless plants can enhance their nutrient acquisition, flowering may therefore impact on future growth and survival.

This study presents evidence for a direct link between reproduction, reflected in a higher relative allocation to inflorescences, turnover and N use-efficiency; the latter being reduced because of increased N turnover. The ranking of species in terms of reproductive allocation was the reverse of their ranking in terms of their N use-efficiency.

Understanding the partitioning of genetic variance in peripheral and central populations may shed more light on the effects of genetic drift and gene flow on population genetic structure and, thereby, improve attempts to conserve genetic diversity. We analysed genetic structure of peripheral and central populations of three insect-pollinated violets (Viola elatior, Viola pumila, Viola stagnina) to evaluate to what extent these patterns can be explained by gene flow and genetic drift. Amplified fragment length polymorphism was used to analyse 930 individuals of 50 populations. Consistent with theoretical predictions, peripheral populations were smaller and more isolated, differentiation was stronger, and genetic diversity and gene flow lower in peripheral populations of V. pumila and V. stagnina. In V. elatior, probably historic fragmentation effects linked to its specific habitat type were superimposed on the plant geographic (peripheral-central) patterns, resulting in lower relative importance of gene flow in central populations. Genetic variation between regions (3-6%), among (30-37%) and within populations (60-64%) was significant. Peripheral populations lacked markers that were rare and localized in central populations. Loss of widespread markers in peripheral V. stagnina populations indicated genetic erosion. Autocorrelation within populations was statistically significant up to a distance of 10-20 m. Higher average genetic similarity in peripheral populations than in central ones indicated higher local gene flow, probably owing to management practices. Peripheral populations contributed significantly to genetic variation and contained unique markers, which made them valuable for the conservation of genetic diversity.

In rare plants that often occur in small or isolated populations the probability of selfing between close relatives is increased as a consequence of demographic stochasticity. The mode of pollination (selfing, outcrossing) may have considerable effects on seed traits and offspring performance and hence potential viability. Since current efforts aiming at the restoration of floodplain grasslands through the transfer of plant material from species-rich source stands may lead to the establishment of initially small populations consisting of founders from different populations, the present paper experimentally investigated the effects of pollen source and floral types (i.e. chasmogamous (CH) and cleistogamous (CL) flowers) on seed traits and offspring performance in three highly endangered violet species (Viola elatior, V. pumila, V. stagnina) of these grasslands. We estimated inbreeding depression and tested the performance of selfed and outcrossed offspring in two microbial environments, i.e. in soil inoculated with (i) non-sterile substrate from the same species ('home'-conditions) and (ii) sterilised substrate. Plants produced more CL capsules than CH flowers. Pollinator exclusion had only small effects on CH seed production. CL seeds had a significantly lower mass per seed than CH seeds. This may be related to constraints in allocation or environmental conditions. Seedling growth was reduced in plants grown under 'home'-conditions as compared to control soils. Under 'home'-conditions, relative fitness of selfed seedlings of V. stagnina was significantly higher than that of crossed progeny. Our results suggest that high genetic differentiation among populations as a consequence of isolation may result in outbreeding depression, e.g., through biochemical or physiological incompatibilities between genes or the breaking of coadapted gene complexes. In V. stagnina, offspring fitness differed considerably between environments, but in general we found no indications for inbreeding depression in these rare species. (c) 2005 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Most studies on the adaptive significance of plant traits compared species that differed in inherent growth rate or species of habitats at the extremes of environmental gradients. In the present paper, we compared the response of Viola elatior and V. pumila, two closely related and morphologically similar violets to the experimental variation of nitrogen availability. The species occur in adjacent habitat types, viz. alluvial forest fringes (V. elatior) and floodplain meadows (V. pumila), that differ in resource availability and disturbance across a relatively small gradient. We tested the hypotheses that, according to their typical habitat, V. elatior should show more traits of a stress tolerant competitive plant, while V. pumila should exhibit more ruderal traits. We carried out a two-year common garden experiment with two levels of fertiliser addition (N5, 5 g N m(-2) yr(-1); N20, 20 g N m(-2) yr(-1)), and analysed a number of traits related to relative growth rate (RGR), allocation, and nutrient use. We further did an experiment with seeds from the experimental plants to study variation in germination rate. We found considerable ontogenetic drift in the species during two years of growth. In the first year, both species showed similar mass and growth rate but these were achieved through different trait combinations. In the second year, V. elatior had a higher RGR, produced more biomass per unit nutrient and developed more leaf area per unit plant mass than V. pumila, while the latter produced thinner leaves. V. pumila had a higher capacity for nutrient acquisition, but a large proportion of the captured nutrients were used for reproduction. Germination rate was higher in V. pumila than in V. elatior, independent of the nutrient status of the mother plant, while seeds of V. elatior from N20 plants had a significantly higher germination rate (67%) than seeds from N5 plants (35%). Our data suggested that many of the analysed trait differences seem to be consistent with the requirements of the different habitat types.

Background and Aims A meta-analysis of global change experiments in arctic tundra sites suggests that plant productivity and the cover of shrubs, grasses and dead plant material (i.e. litter) will increase and the cover of bryophytes will decrease in response to higher air temperatures. However, little is known about which effects these changes in vegetation structure will have on seedling recruitment of species and invasibility of arctic ecosystems. Methods A field experiment was done in a bryophyte-dominated, species-rich subarctic heath by manipulating the cover of bryophytes and litter in a factorial design. Three phases of seedling recruitment (seedling emergence, summer seedling survival, first-year recruitment) of the grass Anthoxanthum alpinum and the shrub Betula nana were analysed after they were sown into the experimental plots. Key Results Bryophyte and litter removal significantly increased seedling emergence of both species but the effects of manipulations of vegetation structure varied strongly for the later phases of recruitment. Summer survival and first-year recruitment were significantly higher in Anthoxanthum. Although bryophyte removal generally increased summer survival and recruitment, seedlings of Betula showed high mortality in early August on plots where bryophytes had been removed. Conclusions Large species-specific variation and significant effects of experimental manipulations on seedling recruitment suggest that changes in vegetation structure as a consequence of global warming will affect the abundance of grasses and shrubs, the species composition and the susceptibility to invasion of subarctic heath vegetation.

Since inference concerning the relative effects of propagule pressure, biotic interactions, site conditions and species traits on the invasibility of plant communities is limited, we carried out a field experiment to study the role of these factors for absolute and relative seedling emergence in three resident and three non-resident confamilial herb species on a nutrient-poor temperate pasture. We set up a factorial field experiment with two levels each of the factors litter cover (0 and 400 g m(-2)), gap size (0.01 and 0.1 m(2)) and propagule pressure (5 and 50 seeds) and documented soil temperature, soil water content and relative light availability. Recruitment was recorded in spring and autumn 2010 and in spring 2011 to cover initial seedling emergence, establishment after summer drought and final establishment after the first winter. Litter alleviated temperature and moisture conditions and had positive effects on proportional and absolute seedling emergence during all phases of recruitment. Large gaps presented competition-free space with high light availability but showed higher temperature amplitudes and lower soil moisture. Proportional and absolute seedling recruitment was significantly higher in large than in small gaps. In contrast, propagule pressure facilitated absolute seedling emergence but had no effects on proportional emergence or the chance for successful colonisation. Despite significantly higher initial seedling emergence of resident than non-resident species, seed mass and other species-specific traits may be better predictors for idiosyncratic variation in seedling establishment than status. Our data support the fluctuating resource hypothesis and demonstrate that the reserve effect of seeds may facilitate seedling emergence. The direct comparison of propagule pressure with other environmental factors showed that propagule pressure affects absolute seedling abundance, which may be crucial for species that depend on other individuals for sexual reproduction. However, propagule batch size did not significantly affect the chance for successful colonisation of disturbed plots.

The invasive herb Lupinus polyphyllus has been focus of a number of fact sheets worldwide but a comprehensive summary of the species’ taxonomy and morphology, distribution, habitat requirements, and biology has been lacking. This paper gives a thorough account of the species’ systematic position and taxonomy, highlighting the difficulties to delimit taxa, which is related to interbreeding among members of this genus. However, L. polyphyllus var. polyphyllus is apparently the taxon that has naturalized and is regionally invasive in temperate-humid climates worldwide. We also present an updated distribution map of L. polyphyllus in the native and invaded ranges, which highlights seven regions in the world where the species has been established. We show that the climatic niche of L. polyphyllus in the invaded range shifts towards higher summer precipitation and lower isothermality, probably because the invaded range includes subcontinental regions of eastern Europe and western Siberia. The habitats of L. polyphyllus range from rather dry to wet, have moderately acidic to strongly acidic soils, and the species’ indicator values across Europe suggest that it occurs along a gradient from very nutrient poor sites to intermediate to rich sites from northern to southern Europe. The species shows high resistance to both drought and frost. In Central Europe, the species has a stronghold in alpic mountain hay meadows, abandoned meadows and pastures, low and medium altitude hay meadows, anthropogenic herb stands and temperate thickets and scrubs. In northern Europe, the species occurs in anthropogenic herb stands along roads and railroads as well as in abandoned pastures and fields. We also found some doubtful information about L. polyphyllus in the literature. This refers to its description as “rhizomatous perennial” although it lacks rhizomes; an apparently very high longevity of its seeds, which may only be true under artificial conditions in an ex situ seed repository; and a very deep rooting depth, which may not represent the average rooting depth but rather an extreme value. Knowledge about the interrelationships between the species’ future population dynamics and spread and ongoing climate warming is lacking. Finally, our review points out that there is currently no evidence-based strategy for a cost-efficient management of L. polyphyllus although it is among the most problematic non-native plant species in Europe due to its environmental and socio-economic impacts. 

There is a growing body of research that recognizes the potentials of biochar application in agricultural production systems. However, little is known about the effects of biochar, especially hydrochar, on production of containerized seedlings under nursery conditions. This study aimed to test the effects of hydrochar application on growth, quality, nutrient and heavy metal contents, and mycorrhizal association of containerized pine seedlings. The hydrochar used in this study was produced through hydrothermal carbonization of paper mill biosludge at 200 °C. Two forms of hydrochar (powder and pellet) were mixed with peat at ratios of 10% and 20% (v/v) under three levels of applied commercial fertilizer (nil, half and full rates). Application of hydrochar had positive or neutral effects on shoot biomass and stem diameter compared with control seedlings (without hydrochar) under tested fertilizer levels. Analysis of the natural logarithmic response ratios (LnRR) of quality index and nutrient and heavy metal uptake revealed that application of 20% (v/v) hydrochar powder or pellet with 50% fertilizer resulted in same quality pine seedlings with similar heavy metal (Cu, Ni, Pb, Zn and Cr) and nutrient (P, K, Ca and Mg) contents as untreated seedlings supplied with 100% fertilizer. Colonization percentage by ectomycorrhizae significantly increased when either forms of hydrochar were applied at a rate of 20% under unfertilized condition. The results of this study implied that application of proper rates of hydrochar from biosludge with adjusted levels of liquid fertilizer may reduce fertilizer requirements in pine nurseries.

Numerous restoration campaigns focused on re-establishing species-rich floodplain meadows of Central Europe, whose species composition is essentially controlled by regular flooding. Climate change predictions expect strong alterations on the discharge regime of Europe's large rivers with little-known consequences on floodplain meadow plants. In this study, we aim to determine the effects of flooding on seedlings of different ages of four typical flood meadow species. To this end, we flooded seedlings of two familial pairs of flood meadow species of wetter and dryer microhabitats for 2 weeks each, starting 2, 4, 6, and 8 weeks after seedling germination, respectively. We show that a 2-week-flooding treatment had a negative effect on performance of seedlings younger than 6 weeks. Summer floods with high floodwater temperatures may have especially detrimental effects on seedlings, which is corroborated by previous findings. As expected, the plants from wet floodplain meadow microhabitats coped better with the flooding treatment than those from dryer microhabitats. In conclusion, our results suggest that restoration measures may perform more successfully if seedlings of restored species are older than the critical age of about 6 weeks before a spring flooding begins. Seasonal flow patterns may influence vegetation dynamics of floodplain meadows and should, therefore, be taken into account when timing future restoration campaigns.

Semi-natural grasslands invaded by the legume Lupinus polyphyllus need the introduction of target species in order to promote highly endangered native target plant communities. However, which techniques are best suited to achieve both invader control and the introduction of target species at the same time? Few studies have investigated restoration techniques that support native plant communities in mountain meadows and control the invader simultaneously. We employed the restoration techniques seed bank activation and green hay transfer in combination with manual removal of the invasive L. polyphyllus on three types of grassland (Nardus grassland, mesic and wet mountain hay meadows) in the low mountain ranges of the Rhon UNESCO Biosphere Reserve in central Germany. Vegetation reacted differently to specific measures depending on the type of grassland. L. polyphyllus cover could be effectively reduced by L. polyphyllus removal in Nardus grassland and mesic hay meadow sites, but not in wet meadows. In Nardus grassland, the cover of target species was lowered by the application of green hay. The target species cover of wet mountain hay meadows declined in response to seed bank activation. When restoring mountain meadows, restoration practitioners should thus first consider the specific vegetation types. As our study indicates that target mountain meadow species may react negatively to restoration measures in the short term, the long-term effects of the application of green hay should be studied.

Plant invasions are among the key drivers of global biodiversity and ecosystem change. They often cause reductions in native species richness and overall biodiversity. Nitrogen-fixing plants are problematic as they affect soil nutrient availability and outcompete species of nutrient-poor sites. Here we assessed the impacts of the legumeLupinus polyphylluson species and functional diversity of mountain meadow communities in the UNESCO Biosphere Reserve Rhon. We compared species diversity (richness, evenness and effective species number), functional diversity (functional richness, evenness, divergence and dispersion) and similarity of plots in three characteristic vegetation types (Nardusgrassland, mesic and wet mountain hay meadows) between different lupine cover classes. We calculated community weighted means (CWMs) of single plant traits and plotted them against lupine cover classes. The invasion ofL. polyphyllushomogenizes vegetation composition since the similarity among plots of the different vegetation types increased with increasing lupine cover. It significantly affected species diversity in terms of richness and effective species number and the functional divergence of the vegetation. The trait set of species occurring together with lupine was shifted towards more competitive trait values. We demonstrate strongly negative impacts ofL. polyphylluson different mountain meadow vegetation types sinceL. polyphyllus, fosters the growth of competitive species and leads to overall more productive plant communities.

Question: We addressed the importance of gut-mediated dispersal by greylag geese for vascular plants in archipelago environments and asked: (a) What proportion of the local species pool is dispersed by geese? (b) Which plant traits characterize species dispersed by geese? (c) Which plant communities are likely to benefit from endozoochory by geese?. Location: Three Swedish Baltic archipelagos. Methods: Goose droppings were collected on 45 islands. Plants germinating from the droppings represent the endozoochorous species pool (ESP). On 108 islands, the presence of vascular plants was recorded in each habitat. These species represent the island species pool (ISP). Differences in functional traits between ESP and ISP were expressed as effect sizes and tested using meta-regressions. Using indicator species analyses and indicator species for managed semi-natural grasslands, we identified the primary habitats of the ESP. Results: Geese dispersed viable diaspores of 97 plant species, which represents 22% of the ISP. Most ESP species were typical for small islands. Geese dispersed a higher proportion of graminoids and less woody plants, higher proportions of chamaephytes and therophytes and less phanerophytes; annuals and bi-annuals were significantly overrepresented. On average, seed volume of the ESP was 95% smaller than that of the ISP. About 51% of all ESP species were dispersed in at least two archipelagos. Geese showed a bias towards species of rocky shore habitats. Conclusion: Geese potentially disperse large amounts of diaspores of many terrestrial island plant species. Through their feeding behaviour, geese select species with certain suites of traits from the regional species pool. Plant dispersal by geese may benefit plants species of rocky shores, but species of formerly managed semi-natural grasslands may also find refuge sites on epilittoral shores after goose-mediated dispersal. The relative importance of geese as dispersal vectors may increase under on-going land-use changes and cessation of grazing networks.

Most island-ecology studies focus on the properties of entire island communities, thus neglecting species-environment relationships operating at the habitat-level. Habitat-specific variation in the strength and sign of these relationships will conceal patterns observed on the island scale and may preclude a mechanistic interpretation of patterns and processes. Habitat-specific species-environment relationships may also depend on the descriptor of ecological communities. This paper presents a comprehensive plot-based analysis of local vegetation composition and species diversity (species richness and species evenness) of (i) rocky shore, (ii) semi-natural grassland and (iii) coniferous forest habitats in three Baltic archipelagos in Sweden. To identify differences and consistencies between habitats and descriptors, we assessed the relative contributions of the variable-sets “region”, “topography”, “soil morphology”, “soil fertility”, “soil water”, “light availability”, “distance” and “island configuration” on local vegetation composition, species richness and species evenness. We quantified the impact of “management history” on the descriptors of local grassland communities by a newly introduced grazing history index (GHI). Unlike species diversity, changes in vegetation composition were related to most of the variable-sets. The relative contributions of the variable-sets were mostly habitat-specific and strongly contingent on the descriptor involved. Within each habitat, richness and evenness were only partly affected by the same variable-sets, and if so, their relative contribution varied between diversity proxies. Across all habitats, soil variable-sets showed highly consistent effects on vegetation composition and species diversity and contributed most to the variance explained. GHI was a powerful predictor, explaining high proportions of variation in all three descriptors of grassland species communities. The proportion of unexplained variance was habitat-specific, possibly reflecting a community maturity gradient. Our results reveal that species richness alone is an incomplete representation of local species diversity. Finally, we stress the need of including habitat-based approaches when analyzing complex species-environment relationships on islands.

Islands are vulnerable ecosystems worldwide, increasingly exposed to human pressure, global climate change and invasive species. Thus, understanding island species diversity is key for nature conservation. Recent studies on insular plant communities indicated that habitat-specific species composition and richness might largely affect diversity patterns observed at the island scale. In consequence, habitat-based approaches are needed to (i) estimate how environmental changes at the habitat scale may affect island diversity, and to (ii) estimate the contribution of different patches of the same habitat to island diversity with respect to habitat-specific environmental constraints. In the present study, we tested these habitat-to-island diversity relationships for shoreline habitats (brackish reeds, salt marsh, rocky shore, tall herbs) and island interior habitats (rocks, semi-natural grassland, pioneer forest, coniferous forest, mixed forest) using 108 islands of three Baltic archipelagos in Sweden. These islands differed in terms of island-scale variables describing effects of island configuration and distance, and habitat-scale variables representing the effects of habitat area, abiotic environment and land-use. The studied habitats differed in their contribution to island species diversity, called habitat specificity. Shoreline habitats shared many common specialist species adapted to extreme conditions like sea salt or bird grazing, while habitats of the island interior harbored mainly species adapted to the specific conditions of a single habitat. We found high variability in habitat specificity as a consequence of habitat-specific environmental factors. Variability was highest for grasslands, where it was related to abandonment and soil fertility, stressing the importance of grassland management for maintaining island biodiversity. Habitats with high habitat specificity through either high species richness or many habitat-specific specialists should be the primary targets for biodiversity management. (C) 2019 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Herbivory through gastropods has among others been proposed as a potential factor responsible for the river corridor distribution of plant species, which is a well known but poorly understood ecological pattern. Since floodplains are characterised by seasonally changing abiotic conditions, viz, floods during winter and spring and severe summer drought that are unsuitable for gastropods they may present safe habitats for highly palatable plant species. In the present study we compared species composition of gastropods and vegetation of twelve grassland sites situated within three floodplain compartments along the Upper Rhine. Additionally, we studied the palatability of 7 days and 25 days old seedlings of five typical floodplain plant species and five mesic grassland species to the slug Deroceras reticulatum in laboratory experiments. Our results showed that both vegetation and gastropod community composition but not gastropod diversity and abundance differed between floodplain compartments. Owing to omnivory of most gastropods the similarity structure of sites based on plants and gastropods was not significantly correlated. In general, slug herbivory significantly reduced survival and biomass of 7 days old seedlings, but responses were species-specific. In contrast, with the exception of Arabis nemorensis, Viola pumila and Taraxacum sect. Ruderalia biomass of 25 days old seedlings was not significantly affected by slug herbivory. Although the response of floodplain plant species as a group to slug herbivory did not differ from common grassland species, our results suggest that gastropods may potentially influence the distribution pattern of the highly palatable river corridor species Arabis nemorensis and Viola pumila. However, further research is needed to estimate the damage to river corridor plants through gastropod herbivory and its effect on competitive relationships under natural conditions.

Riparian zones are species-rich and functionally important ecotones that sustain physical, chemical and ecological balance of ecosystems. While scientific, governmental and public attention for riparian zones has increased over the past decades, knowledge on the effects of the majority of anthropogenic disturbances is still lacking. Given the increasing expansion and intensity of these disturbances, the need to understand simultaneously occurring pressures grows. We have conducted a literature review on the potential effects of anthropogenic pressures on boreal riparian zones and the main processes that shape their vegetation composition. We visualised the observed and potential consequences of flow regulation for hydropower generation, flow regulation through channelisation, the climate crisis, forestry, land use change and non-native species in a conceptual model. The model shows how these pressures change different aspects of the flow regime and plant habitats, and we describe how these changes affect the extent of the riparian zone and dispersal, germination, growth and competition of plants. Main consequences of the pressures we studied are the decrease of the extent of the riparian zone and a poorer state of the area that remains. This already results in a loss of riparian plant species and riparian functionality, and thus also threatens aquatic systems and the organisms that depend on them. We also found that the impact of a pressure does not linearly reflect its degree of ubiquity and the scale on which it operates. Hydropower and the climate crisis stand out as major threats to boreal riparian zones and will continue to be so if no appropriate measures are taken. Other pressures, such as forestry and different types of land uses, can have severe effects but have more local and regional consequences. Many pressures, such as non-native species and the climate crisis, interact with each other and can limit or, more often, amplify each other's effects. However, we found that there are very few studies that describe the effects of simultaneously occurring and, thus, potentially interacting pressures. While our model shows where they may interact, the extent of the interactions thus remains largely unknown.

Question

Riparian vegetation and seed banks are the foundation of functional riparian zones, yet insight in the processes that explain their composition is limited. We tested three theories fundamental to dispersal and environmental filtering of riparian seed banks and vegetation. Combining these theories, we expected hydrochory to lead to increased species richness downstream in both soil seed bank and vegetation with accumulation restarting after a lake section, and geomorphological filtering to lead to higher similarity of seed bank and vegetation composition locally and within lakes, slow-flowing sections and rapids, respectively, than between them.

Location

Svartån, a free-flowing river in central Sweden.

Methods

We surveyed riparian vegetation and conducted a germination experiment on riparian soil seed bank cores from lakes, slow-flowing sections and rapids. We combined these with trait data on seed dispersal syndromes, floating capacity of seeds and seed longevity. We analysed differences throughout the system with linear models and between process domains with Kruskal–Wallis tests and similarity in community composition with the Horn–Morisita similarity index.

Results

Our results indicated that species richness did not increase downstream and was relatively similar throughout lakes, slow-flowing sections and rapids for both riparian seed banks and vegetation. Seed floating capacity was similar throughout these river sections, too, and seed longevity was higher in seed banks than in vegetation.

Conclusions

Geomorphology and hydrochory were not as important drivers in this area for riparian seed bank and vegetation composition as expected. In the area and on the scale studied here, other local and regional environmental factors and dispersal syndromes are more likely to be determinants of riparian dynamics and composition. Continued studies of riparian seed banks can help improve our understanding of riparian composition and functioning in the future.

Premise of research. Halophyte species grow where salt concentrations are high. Still, their germination may be affected by salts, either by creating an osmotic potential that prevents water uptake or by dissociating in ions that can cause different grades of toxicity. With the increase of salinized areas, it becomes important to understand the behavior of these species. Methodology. We studied how the germination of five halophyte species that occur along a salinity gradient in the Bahia Blanca coastal zone, Atriplex undulata, Cyclolepis genistoides, Allenrolfea patagonica, Sarcocornia perennis, and Heterostachys ritteriana, responds to variations in osmotic agents, osmotic potential, and temperature. Seeds were exposed to different osmotic potentials using NaCl (neutral salt), Na2CO3 (alkaline salt), and mannitol solutions in a germination chamber experiment. Germination was recorded during 42 d. Germination percentage, mean germination time, and synchrony were calculated. Pivotal results. Our experimental results showed that for the five halophyte species under study, germination was mostly driven by osmotic potentials and osmotic agents. At high osmotic potential, the germination response did not differ significantly from controls, except for Allenrolfea and Cyclolepis, which showed lower germination when treated with Na2CO3. Low osmotic potentials and Na2CO3 were detrimental to germination, reflected by lower germination percentages, higher mean germination times, and lower synchrony. Conclusions. In general, the response to the alkaline salt was more negative than that to the neutral salt or mannitol, regardless of the species. Each species showed a different response to the salts under study, and this response matched well with the distribution of species along the salinity gradient observed in the field.

Jung LS, Eckstein RL, Otte A & Donath TW (2012). Above- and below-ground nutrient and alkaloid dynamics in Colchicum autumnale: indications for optimal mowing dates for population control or low hay toxicity. Weed Research52, 348-357. Summary In some Central European regions, the conservation of seminatural grasslands is jeopardised by management intensification or abandonment, caused by high densities of the toxic weed Colchicum autumnale. We investigated two possibilities to deal with C. autumnale: (i) reducing population densities by mowing when nutrient contents are high in leaves and capsules and low in the storage organs, that is, when the ratio between both is highest, or (ii) reducing alkaloid content in hay by mowing when alkaloid content of leaves and capsules of C. autumnale is low. To identify the optimal mowing point, we analysed the dynamics of nutrients, starch and alkaloids of naturally grown plants in two biogeographical regions. In the colder region, the maximum nutrient ratio between above-ground and storage organs, as well as alkaloid content in leaves and capsules, occurred significantly later. Compared with the common first mowing date (15 June), alkaloid content decreased significantly until 5 July in both regions. On both dates, it was on average 1.8 times higher in the colder region. Our results suggest the following time for the two management options: (i) mowing at about 25 cm plant height (late April/early May) to reduce C. autumnale densities or (ii) delayed mowing in late June/early July when the plant has turned brown and dry and alkaloid content has declined.

Colchicum autumnale is a perennial hysteranthous geophyte, which is native to Europe. It is characteristic of periodically wet to moderately moist grasslands but also occurs in alluvial forests. This article gives an overview of the taxonomy, distribution, life cycle, and population biology of C. autumnale and puts special emphasis on its morphology, germination and its response to competition and management.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

Aim We assessed the role of mowing machinery and endozoochory by migratory sheep as dispersal vectors in semi-natural grasslands by comparing the species compositions and traits of species found in the vectors to the regional above-ground vegetation and soil seed bank. Furthermore, we discuss how their interplay may affect the conservation of semi-natural grasslands. Location Rhon Mountains, central Europe. Methods Plant material from mowers (n = 12 from one date) and dung samples from migratory sheep (n = 39 from 13 dates) were collected and the dispersed plant species were determined using the emergence method. We compared the species compositions to the regional above-ground vegetation and seed bank using non-metric multidimensional scaling (NMDS) and indicator species analysis. Furthermore, we compared functional traits of the dispersed species to traits of non-dispersed species of the regional species pools by calculating log-response ratios and performing metaregressions. Results While 43 species were shared between the vectors, the vegetation compositions differed from each other. Mower samples were more similar to the above-ground vegetation whereas dung samples were more similar to the seed bank. Mowers and sheep endozoochory favoured the dispersal of species with different traits and phenologies. Species with small seed sizes were prevalent in both vectors. Mowers were less selective concerning most traits, but favoured high-growing grasses such as Alopecurus pratensis and Trisetum flavescens. Sheep dung samples contained less grasses and more palatable species, such as Urtica dioica. Mowers were most selective concerning phenology, whereas endozoochory by migratory sheep also included late-flowering species. Conclusion Sheep endozoochory and mowing machinery are complementary dispersal vectors favouring species with differing functional traits. Sheep endozoochory enables dispersal of species that have unfavourable traits (e.g. low releasing heights) or phenologies for dispersal by mowing machinery. To ensure the dispersal of a high number of plant species in semi-natural grasslands, the interplay of different vectors should be considered.

In semi-natural grasslands, mowing leads to the dispersal of species that have viable seeds at the right time. For invasive plant species in grasslands, dispersal by mowing should be avoided, and information on the effect of cutting date on the germination of invasive specks is needed. We investigated the germination of seeds of the invasive legume Lupinus polyphyllus Lind!. depending on the cutting date. We measured seed traits associated with successful germination that can be assessed by managers for an improved timing of control measures. To this end, we sampled seeds of L polyphyllus on six cutting dates and analyzed the germination of these seeds in climate chambers and under ambient weather conditions. We collected information on seed morphology (color/size/hardseededness) for each cutting date to identify seed traits associated with successful germination. Observed germination patterns were highly asynchronous and differed between seeds cut at different dates. Seeds cut early, being green and soft, tended to germinate in autumn. Seeds cut late, being dark and hard, were more prone to germinate the following spring, after winter stratification. This allows the species to utilize germination niches throughout the year, thus indicating a bet-hedging strategy. Seed color and the percentage of hard seeds were good predictors of germination percentage, but not of mean germination time and synchrony. Managers should prevent the species producing black and hard seeds, while cutting plants carrying green and soft seeds is less problematic. Furthermore, germination patterns differed between climate chambers and the common garden, mainly because germination of dormant seeds was lower in climate chambers. More germination experiments under ambient weather conditions should be carried out, as they can give information on the germination dynamics of invasive species.

Photo observations are a highly valuable but rarely used source of citizen science (CS) data. Recently, the number of publicly available photo observations has increased strongly, for example, due to the use of smartphone applications for species identification. This has enabled the raising of ecological insights in poorly studied subjects. One of the fields with the highest potential to benefit from the use of photo observations is phenology. We propose a workflow for iPhenology, the use of publicly available photo observations to track phenological events at large scales. The workflow comprises data acquisition, cleaning of observations, phenological classification and modelling spatiotemporal patterns of phenology. We explore the suitability of iPhenology to observe key phenological stages in the plant reproductive cycle of a model species and discuss limitations and future prospects of the approach using the example of an invasive species in Europe. We show that iPhenology is suitable to track key phenological events of widespread species. However, the number and quality of available observations may differ among species and phenological stages. Overall, publicly available CS photo observations are suitable to track key phenological events and can thus significantly advance the knowledge on the timing and drivers of plant phenology. In future, integrating the workflow with automated image processing and analysis may enable real-time tracking of plant phenology.