Abandonment of traditionally managed mesic mountain meadows affects plant species composition and diversity

In recent years abandonment of traditional management of mountain grasslands has been observed throughout Central Europe. However, the impact of abandonment on vegetation of mountain grasslands is still unclear. In this study it was hypothesized that the cessation of traditional management of mesic mountain meadows causes changes in their species composition and a decrease in the biodiversity. In total, 260 plots were established in the Sudetes (SW Poland) on meadows with regular annual mowing, meadows with irregular mowing management, and abandoned meadows. Relevés (5 × 5 m) were performed, and the habitat properties were determined using Ellenberg indicator values. The study confirmed the hypothesis that the various ways of extensive management have an influence on species richness. The lowest species richness was observed on the irregularly managed meadows, while higher species numbers were found on the abandoned and regular managed meadows. The majority of patches on abandoned meadows exhibited degradation through the expansion of Solidago gigantea, Solidago canadensis, Lupinus polyphyllus, Heracleum sosnovsky, Calamagrostis epigejos, Deschampsia flexuosa, Festuca rubra and Hypericum maculatum. Meadows subjected to different management practices differed significantly in Ellenberg indicator values. The abandoned meadows had the highest values of the light index (L) and nitrogen availability (N), whereas the highest values of soil moisture (F) were noted on the irregularly managed meadows. The degradation of mountain mesic meadows requires regular mowing management, which stops ecological succession and preserves their high biodiversity.     

Deceptive pollination of Dactylorhiza incarnata: an experimental test of the magnet species hypothesis

Floral deception, which mainly appears in highly evolved families such as Orchidaceae, was studied in Central Finland. In nectarless Dactylorhiza incarnata, the deceptive pollination system has been considered to function best in remote habitats such as marshes, where flowering plants attractive to pollinators are rare (remote habitats hypothesis). In contrast, the magnet-species theory predicts that a nectarless plant benefits from growing in the vicinity of nectarcontaining species. We tested these hypotheses by adding attractive, nectar-containg violets (Viola x wittrockiana) to orchid populations. The percentage of fruit set in D. incarnata was adversely affected by the violets, probably because interspecific exploitation competition for pollinators took place in favour of the violas at the expense of the orchids. This result gave no support for the magnet-species theory and supported the remote habitats hypothesis.     

Differential and reversible responses of common fen meadow species to abandonment

Abstract. We studied the effects of abandonment on two common fen plant species. In mown and a chronosequence of abandoned fen meadows spanning 35 yr, we measured fitness traits of the sedge Carex davalliana and the forb Succisa pratensis. Cessation of mowing had little effect on fitness traits and seed production of C. davalliana, but seedling density decreased more than threefold. Population density of S. pratensis decreased with increasing community biomass, but was not affected by the cessation of mowing. However, flowering frequency increased threefold and seed production was 20% higher in fallow meadows. Consequently, seedling density of S. pratensis increased nearly threefold after abandonment. However, these changes were not dependent on the age of the fallow. In a common garden and germination experiment, we found no differences in either species between plants from fallows and mown fen meadows, except for the height of the flowering stalk of S. pratensis. The combined results from the common garden experiment and the field studies indicate that changes in fitness traits observed in fallows were mostly phenotypic and likely to be reversible. If other species react in similar ways, there is a high potential for re‐establishing traditional fen meadow communities from fallows by mowing.     

Polymorphic populations of Dactylorhiza incarnata s.l. (Orchidaceae) on the Baltic island of Gotland: morphology, habitat preference and genetic differentiation

Background and Aims

Organisms may be polymorphic within natural populations, but often the significance and genetic background to such polymorphism is not known. To understand the colour polymorphism expressed in the diploid marsh-orchids Dactylorhiza incarnata, morphological, habitat and genetic differentiation was studied in mixed populations on the island of Gotland, supplemented with genetic marker data from adjacent areas.

Methods

A total of 398 accessions was investigated for plastid haplotype and three nuclear microsatellites. Morphometric data and vegetation data were obtained from a subset of 104 plants.

Key Results

No clear pattern of habitat differentiation was found among the colour morphs. Within sites, the yellow-flowered morph (ochroleuca) was slightly larger than the others in some flower characters, whereas the purple-flowered morph with spotted leaves (cruenta) was on average smaller. However, populations of the same colour morph differed considerably between sites, and there was also considerable overlap between morphs. Morphs were often genetically differentiated but imperfectly separated within sites. Most populations were characterized by significant levels of inbreeding. The ochroleuca morph constitutes a coherent, highly homozygous sublineage, although introgression from purple-flowered morphs occurs at some sites. The cruenta morph was genetically variable, although Gotland populations formed a coherent group. Purple-flowered plants with unspotted leaves (incarnata in the strict sense) were even more variable and spanned the entire genetic diversity seen in the other morphs.

Conclusions

Colour polymorphism in D. incarnata is maintained by inbreeding, but possibly also by other ecological factors. The yellow-flowered morph may best be recognized as a variety of D. incarnata, var. ochroleuca, and the lack of anthocyanins is probably due to a particular recessive allele in homozygous form. Presence of spotted leaves is an uncertain taxonomic character, and genetic differentiation within D. incarnata would be better described by other morphological characters such as leaf shape and stature and size and shape of lip and spur.Key words: Dactylorhiza incarnata, cruenta, ecology, genetic differentiation, Gotland, microsatellites, ochroleuca, plastid DNA, polymorphism     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.     

Recolonization after habitat restoration leads to decreased genetic variation in populations of a terrestrial orchid

Colonization is crucial to habitat restoration projects that rely on the spontaneous regeneration of the original vegetation. However, as a previously declining plant species spreads again, the likelihood of founder effects increases through recurrent population founding and associated serial bottlenecks. We related Amplified Fragment Length Polymorphism markers genetic variation and fitness to colonization history for all extant populations of the outcrossing terrestrial orchid Dactylorhiza incarnata in an isolated coastal dune complex. Around 1970, D. incarnata suffered a severe bottleneck yet ultimately persisted and gradually spread throughout the spatially segregated dune slacks, aided by the restoration of an open vegetation. Genetic assignment demonstrated dispersal to vacant sites from few nearby extant populations and very limited inflow from outside the spatially isolated reserve. Results further indicated that recurrent founding from few local sources resulted in the loss of genetic diversity and promoted genetic divergence (F_ST = 0.35) among populations, but did not influence population fitness. The few source populations initially available and the limited inflow of genes from outside the study reserve, as a consequence of habitat degradation and spatial isolation, may have magnified the genetic effects of recurrent population founding.     

Systematics of the Dactylorhiza euxina/incarnata/maculata polyploid complex (Orchidaceae) in Turkey: evidence from allozyme data

 Material of Dactylorhiza were sampled from 49 localities in Turkey and investigated for allozyme variation at ten loci (nine enzyme systems). Among diploids, the Anatolian D. osmanica and D. umbrosa were allozymically variable, but not distinct from each other or from D. incarnata. Dactylorhiza saccifera contained the same alleles as the European D. fuchsii. Dactylorhiza iberica and D. euxina were distinct from each other and the other diploids. On basis of allozyme patterns three distinct allotetraploid genotypes were distinguished, and each of them could be treated as a separate species. Dactylorhiza nieschalkiorum is similar to European allotetraploids, and may have arisen from hybridization between D. incarnata s.l. and D. saccifera. Dactylorhiza urvilleana may have arisen from parents related to present-day D. saccifera and D. euxina, but it also contains additional alleles that have not been found in any of the diploids investigated. A third allotetraploid known from four populations in the Ardahan and Kars provinces of north-eastern Turkey combines the allozyme patterns found in material of D. incarnata s.l. from the same area with those from D. euxina. It is here described for the first time as D. armeniaca. Received November 14, 2000 Accepted June 20, 2001     

Nutrient availability and limitation in traditionally mown and in abandoned limestone grasslands: a bioassay experiment

Management is necessary for the conservation of limestone grasslands. However, the traditional management of hay-making every year in July is no longer profitable for farmers. Hence many species-rich grasslands have become abandoned. The aim of this study is (a) to investigate the consequences of abandonment (as compared with annual mowing) on the availability of soil nutrients, and (b) to determine the type of nutrient limitation. The soil was taken from a long-term experimental site set up 22 years ago in northern Switzerland. The availability of soil N and P was assessed in a bioassay where soil from mown and unmown plots was supplied with different nutrient solutions or deionised water as control. Seedlings of Galium mollugo s.str. L. and Raphanus sativus ssp. oleiferus (DC) Metzg. were used as phytometer plants. Their growth in the control treatment was limited by N and P almost to the same extent, indicating that the availability of both elements was very low in this soil. P limited plant growth slightly more, but was overcome in the case of Raphanus by a high P content in the seeds. The N and P availability responded differently to management. The availability of N was slightly higher in soil from the abandoned plots, whereas the P availability did not differ significantly. Accumulation of nitrogen in the soil after abandonment did presumably not have any decisive effect on the vegetation because the amount was small and because the vegetation seemed primarily P-limited. The direct effect of mowing or abandonment on plants is therefore likely to be much more important for species composition than the minor changes in soil nutrients.     

Differential effects of abandonment on the demography of the grassland perennial Succisa pratensis

Abandonment of traditional land-use practices can have strong effects on the abundance of species occurring in agricultural landscapes. However, the precise mechanisms by which individual performance and population dynamics are affected are still poorly understood. To assess how abandonment affects population dynamics of Succisa pratensis we used data from a 4-year field study in both abandoned and traditionally grazed areas in moist and mesic habitats to parameterize integral projection models. Abandoned populations had a lower long-term stochastic population growth rate (λ _S = 0.90) than traditionally managed populations (λ _S = 1.08), while λ _S did not differ between habitat types. The effect of abandonment differed significantly between years and had opposed effects on different vital rates. Individuals in abandoned populations experienced higher mortality rates and lower seedling establishment, but had higher growth rates and produced more flower heads per plant. Population viability analyses, based on a population survey of the whole study area in combination with our demographic models, showed that 32 % of the populations face a high risk of extinction (>80 %) within 20 years. These results suggest that immediate changes in management are needed to avoid extinctions and further declines in population sizes. Stochastic elasticity analyses and stochastic life table response experiments indicated that management strategies would be most effective if they increase survival of small plants as well as seedling establishment, while maintaining a high seed production. This may be achieved by varying the grazing intensity between years or excluding grazers when plants are flowering.     

Assessing the impact of land abandonment, nitrogen enrichment and fairy-ring fungi on plant diversity of Mediterranean grasslands

Semi-natural calcareous grasslands are of great conservation interest because of their high species richness, but they are threatened by land abandonment and nitrogen eutrophication. These plant communities evolved as a result of a long history of human activity, which generated and maintained these habitats by extensive grazing and mowing. Calcareous grasslands are listed as a priority for conservation in the EC Habitats Directive. However, the effects of different management regimes, nitrogen enrichment, and soil-borne pathogens on plant species diversity are less clear for grasslands of the Mediterranean Basin, compared to meadows in Northern and Central Europe. In this study, we assessed the impact of land abandonment, nitrogen enrichment, and fairy-ring fungi on species diversity in semi-natural grasslands found in the Mediterranean Basin by comparing the available literature with findings from recent studies carried out in Central Italy. In a series of field experiments, the cutting of abandoned grassland consistently reduced the living biomass of the dominant perennial grasses, such as Brachypodium rupestre and Bromus erectus, and promoted a rapid increase in species richness and diversity by allowing the establishment of rare species. There was a similar, but less effective, restoration of species diversity and composition in mowed grassland after litter removal. We also show that nitrogen enrichment at levels comparable to atmospheric deposition depresses species diversity, which also hampers the positive effects of litter removal. Our findings are consistent with previous results achieved in Northern and Central Europe, which however, mainly focused on grasslands with intermediate to high primary productivity levels. The limited availability of data from low-productivity, drought-prone Mediterranean grasslands requires further studies to assess the impact of land abandonment and nitrogen eutrophication in such ecosystems. Finally, we discuss the role of fairy-ring fungi in the maintenance of plant diversity in species-rich grassland. We show that fairy-ring fungi (e.g. Agaricus campestris) critically affect the spatial distribution and diversity of coexisting plant species. By killing the dominant perennial herbs, these radially growing plant pathogens produce empty niches for rare, short-lived species, thus affecting the vegetation pattern. Overall, our results are of interest for environmental managers, as they provide guidelines for the restoration of abandoned areas and the conservation of these species-rich habitats.     

Reinstatement of traditional mowing regimes counteracts population senescence in the rare perennial Primula vulgaris

Question: Traditional management of grassland verges or ditch banks included mowing as a way to provide additional harvesting of hay. Nowadays, such sites are often left unmanaged, as mowing verges is no longer profitable in modern agricultural systems. Are vulnerable plant species able to withstand competition with the surrounding vegetation and maintain viable populations under these circumstances? How do they respond to reinstatement of traditional mowing regimes? Location: Oedelem, northwestern Belgium. Methods: To investigate the effect of reinstatement of the rare perennial Primula vulgaris, demography and adult plant performance were monitored in a grassland verge between 1999 and 2003 under different mowing regimes. Year transitions between life stages were analysed with matrix population models. To disentangle the contributions of the deviations in different life stage transitions to the variation in overall population growth rate, life table response experiments were used. Results: Both management and year had a strong impact on demographic traits of P. vulgaris. If plots were left unmanaged, lower plant performance and declining population growth rates were observed. While population growth rates differed significantly between mowing regimes, mowing of plots only in July did not differ from mowing in July and October in terms of vegetative and reproductive output of adults. Mowing twice a year appeared to be most efficient in increasing population growth rate both by raising recruitment and growth of individuals into large reproductive adults. Conclusions: Large P. vulgaris populations show a good ability to recover from recent abandonment of traditional management regimes. By mowing twice a year, managers are able to target vital rates that are most influential: growth and flowering of adult individuals.     

The effects of mulching and abandonment on the viability of the perennial grassland species Plantago lanceolata

Abandonment and fragmentation are the primary causes of decline for many light-demanding grassland species in the modern agricultural landscape. In order to maintain a flora rich in such species, design of management practices based upon knowledge of life-history consequences following management shifts is essential. In this study, we investigated the demography of the perennial grassland species Plantago lanceolata for 4 years in one mulched site and two abandoned sites to compare how the two treatments influenced population dynamics. Compared to abandonment, mulching had a small but positive effect on the population growth rate (λ) of P. lanceolata. In both the mulched and the abandoned sites, growth and stasis among the largest plants contributed most strongly to λ, while the contribution of fecundity was negligible in both treatments. λ of P. lanceolata varied more between years than between treatments, most notably because λ values below unity were observed in the abandoned sites in a 1-year period that included a severe winter. The population of the mulched site consisted of larger ramets and had higher clonal growth rates than the populations of the abandoned sites. This explains the apparently higher resilience to unfavorable climatic conditions in the mulched site. We conclude that management by mulching is better than abandonment for the long-term persistence of P. lanceolata. However, due to low recruitment also in the mulched site, even mulching may be insufficient for P. lanceolata to persist over long time periods.     

Genetic structure of populations and natural hybridization between <Emphasis Type="Italic">Dactylorhiza salina</Emphasis> and <Emphasis Type="Italic">D. incarnata</Emphasis> (Orchidaceae)

The results of studying the polymorphism and genetic structure of populations of D. salina and D. incarnata growing in Zabaykalsky krai and Buryatia are represented according to the data of allozyme analysis of eight genetic loci (PGI, NADHD, SKDH, GDH, PGM, DIA, ADH, and IDH). The specificity of the allelic structure of loci SKDH, PGM, and IDH is established, for which D. salina and D. incarnata reliably differ from each other. It is shown that interspecies introgressive hybrid complexes with different genetic structures were formed in Transbaikalia. Places of mass growth of D. incarnata were observed to have single plants of D. salina, the interspecies hybrids of the first and subsequent generations. Places of mass growth of D. salina were observed to contain only the hybrids that are not hybrids of the first generation. They were heterozygous not for three loci with differentiating alleles of both parents, SKDH, PGM, and IDH, but for only one of them. The degree of genetic differentiation among five populations of D. salina was on average 7.5% and that of D. incarnata was 7.1%, which in accordance with Wright’s estimation relates to mean values. The average value of FST for all studied populations of the two related species of the genus Dactylorhiza was 0.478, indicating a very high degree of genetic differentiation between D. salina and D. incarnata growing in Transbaikalia. The greatest differences between the species are for the allelic structure of loci SKDH, PGM, and IDH (FST was equal to 0.705, 0.976, and 0.762, respectively). Analysis of molecular variance (AMOVA) showed that populations of D. salina and D. incarnata in the zone where their ranges in Zabaykalsky krai and Buryatya overlap have essential differences both for the variation of alleles frequencies of eight loci (71%, d.f. = 9) and for the variability of genotypes (61%, d.f. = 9). Despite the fact that D. salina and D. incarnata explicitly share a gene flow as a result of interspecies hybridization, the genetic differentiation of populations of these related species remains at a high level.     

Comparison of reproductive success in two orchids: the nectarless Dactylorhiza majalis s.s. and the nectar-producing Gymnadenia conopsea s.l.

Differences in reproductive biology between two orchid species were examined: Eight Danish populations of the nectarless Dactylorhiza majalis ssp. majalis and six Swedish populations of the nectar-producing Gymnadenia conopsea ssp. densiflora . Population size varied from 24 to 1700 flowering individuals in D. majalis and from 100 to 1200 in G. conopsea. Dactylorhiza majalis had only half as many flowers per spike as G. conopsea . Fruit-set of D. majalis ranged from 16 to 39%, much lower than the fruit-set of G. conopsea (78–91%). Thus in the species with a few flowers and no nectar a lower fruit-set was observed than in the species with many flowers and presence of nectar. The low fruit-set of D. majalis was pollen-limited and fruit-set was positively correlated with pollinia removal and the latter was negatively correlated with population size. In both species, fruit-set and population size were uncorrelated.     

Plant species diversity in an old, traditionally managed hay meadow compared to abandoned hay meadows in southwest Norway

A chronosequence, representing a successional series, was used for the comparison of a hay meadow site managed in an old traditional way for at least a hundred years, and hay meadow sites abandoned for about 10, 20 and 30 years, respectively. Old traditional management included grazing early and late in the growing season, mowing in August and light or no fertilizing. The tree cover was the most important factor deciding the composition of vegetation. Time since abandonment was not completely correlated to tree cover, as some plots had a dense canopy and others were situated in the open. The total species number decreased with number of years since abandonment in plots > 0.001 m²and <100 m². The highest species number in 1 m² plots was recorded in the managed site, with 38 species of phanerogams. Fourtyeight % of the indicators of traditional management present in the managed site was recorded in the site which had been abandoned for 30 years. Frequency — log area curves made it possible to group species according to persistence in the sward. As a result, a group of functional indicators of rare hay meadows in the region was distinguished.     

Morphology and phenology of Scabiosa columbaria from mown and grazed habitats - Results of a simulation experiment

We analysed the morphology and phenology of Scabiosa columbaria in relation to grassland management by mowing and grazing. We selected 12 populations from mown and grazed calcareous grassland in six regions of central Europe. At each population site, we collected seed material to produce plants for a morphological and phenological analysis of the species in a simulation experiment, which comprised three treatments (control, simulated mowing and simulated grazing).The simulation experiment revealed a clear morphological and phenological differentiation of S. columbaria. Plants from the two treatment types differed in size, since individuals from mown sites had longer top peduncles than individuals from grazed sites. Individuals from mown populations flowered earlier than individuals from grazed populations. In the control treatment, individuals from both habitat types produced a similar number of fruiting capitulums. When subjected to simulated mowing, the reproductive success of the individuals was clearly reduced. Plants from mown habitats produced, however, a significantly larger number of fruiting capitulums than plants from grazed habitats. Simulated grazing reduced the reproductive success stronger than simulated mowing but differences between individuals from both habitat types were not significant.The results of our simulation experiment clearly indicate morphological and phenological adaptation of S. columbaria to mowing and grazing. The comparatively young selection pressure of mowing obviously selected early flowering individuals, while the original land use by grazing favoured late flowering plants.     

The effect of management on productivity,litter accumulation and seedling recruitment in a Carpathian mountain grassland

The management regime may have a significant impact on the productivity and dynamics of grasslands, but the causal relationships influencing grassland conservation value are still not completely understood. Changes of selected community characteristics, such as standing crop, proportion of forbs in the standing crop, litter amount, litter decomposition and seedling recruitment, were investigated in a 4 year manipulative experiment in a mountain grassland in Slovakia. The aim of the research was to compare changes in newly abandoned sites and sites where restoration measures were applied after 20 years of abandonment. The sites were located in areas containing two vegetation types of the Arrhenatherion alliance (wet Poo-Trisetetum and dry Anthoxantho–Agrostietum) with different moisture regimes. The expected increase of the standing crop after abandonment was rather slow, and more pronounced towards the end of the experiment, and in the wet meadow type (~30% increase). The restoration mowing promoted forb proportions in the biomass, but it did not decrease the standing biomass in the restored grasslands. Strong litter accumulation after abandonment was observed in subsequent years after abandonment, when the amount of litter increased about 100% in abandoned plots. Decrease in litter was also significant after the start of restoration mowing (a decrease from 258 to 159 g m⁻² in wet type and from 287 to 147 g m⁻² in dry type was noted). Accumulated litter was negatively correlated to seedling recruitment (r = −0.63 at the end of the experiment). The litterbag experiment showed that the wet type has a higher rate of decomposition, with 20% more biomass decomposed during the litter-bag experiment. The experiment confirmed a negative role of litter accumulation on seedling recruitment, with the number of seedlings per m² decreasing from 413 to 321 individuals in the abandoned wet-type site. This may lead to a decrease in species richness. Mowing along with raking of mowed biomass may be a useful tool to restore degraded mountain grasslands and to remove accumulated litter from the stands.     

Systematics and conservation genetics of Dactylorhiza majalis ssp. elatior (Orchidaceae) on Gotland

A tall allotetraploid member of the Dactylorhiza incarnata/maculata complex with unspotted leaves and large pinkish flowers from the island of Gotland in the Baltic was examined for molecular variation patterns at five nuclear microsatellite loci, nuclear ITS and in plastid haplotypes. The allotetraploid was well separated from allopatric allotetraploids of similar appearance, including the western European D. majalis ssp. integrata (syn. D. praetermissa) and forms of D. majalis ssp. lapponica from mainland Sweden. It also differed from other allotetraploids distributed in the Baltic Sea region, including D. majalis ssp. baltica and D. majalis ssp. lapponica. It is here recognized as D. majalis ssp. elatior (Fr.) Hedrén & H. A. Pedersen. Dactylorhiza osiliensis Pikner, described from Saaremaa (Estonia) is regarded as a synonym. The distribution covers Gotland, Saaremaa and possibly Hiiumaa. Dactylorhiza majalis ssp. elatior may have one or several recent origins within its present distribution area, and it contains no other molecular markers than those found in the parental D. incarnata var. incarnata and D. maculata ssp. fuchsii in the same area. It appears to have weak barriers towards secondary hybridization with its parental lineages. The situation is reminiscent to that of other young allotetraploids in the D. majalis s.l. complex, suggesting that introgression may be an underestimated process explaining the accumulation of genetic diversity in evolving allopolyploid plants.     

Does Germination Success Differ with Respect to Seed Mass and Germination Season? Experimental Testing of Plant Functional Trait Responses to Grassland Management

BACKGROUND AND AIMS: Plant functional trait responses to processes such as grassland management have been analysed frequently; however, the scaling-up from individual traits to the outcomes of vegetation dynamics has seldom been tested. In this experiment, germination success was studied with respect to the relationships between grassland management (mowing and grazing), as well as abandonment, and two traits that are relevant for seedling recruitment: seed mass and germination season. On the basis of discussions in the literature and indirect trait analyses in our previous studies, the following hypotheses are proposed: (1) with respect to seed mass, mowing and grazing favour the germination of small seeds, whereas after abandonment the germination success of larger seeds is higher; and (2) with respect to germination season, mowing and grazing favour autumn-germinating seeds, whereas succession promotes spring-germinating seeds. METHODS: The germination experiment took place in a semi-natural, dry grassland in north-east Germany. Seeds of eight herbaceous species that differ with respect to seed mass and germination season were sown in mown, grazed and abandoned plots. Germination success was documented during the following year. KEY RESULTS AND CONCLUSIONS: Contrary to the hypothesis, germination of small seeds was not promoted by mowing or grazing and they germinated relatively more often than expected in the abandoned plots. A relationship between abandonment and gaps of bare soil below the vegetation cover that favour germination of small seeds was likely, but could not be proved statistically. It is possible that the small seeds suffered less from predation. Mowing favoured autumn germination, which could be explained by the removal of biomass in late summer. Contrary to our expectation, there was relatively more spring germination after grazing than after mowing, yet vegetation height was smallest in spring. Generally, germination season was found to be related to the temporal occurrence of favourable light conditions.