Summary1. Plant litter is a key component in terrestrial ecosystems. It plays a major role in nutrient cycles and community organization. Land use and climate change may change the accumulation of litter in herbaceous ecosystems and affect plant community dynamics. Additionally, the transfer of seeds containing plant material (i.e. litter) is a widespread technique in grassland restoration. 2. Ecosystem responses to litter represent the outcome of interactions, whose sign and strength will depend on many variables (e.g. litter amount, seed size). A previous meta-analysis (from 1999) reported that litter had an overall negative effect on seed germination and seedling establishment in different ecosystems. However, recent studies indicated that this might not be the case in grassland ecosystems. 3. We used 914 data from 46 independent studies to analyse the effects of litter on seedling (i) emergence, (ii) survival and (iii) biomass, employing meta-analytical techniques. Each data set was stratified according to methodology, grassland type, irrigation conditions, litter amount and seed size. 4. We found an overall neutral effect of litter presence on seedling emergence and survival and a positive effect on seedling biomass. However, whereas for field experiments the response remained neutral, it was positive for common garden studies. In glasshouse experiments, litter effects were negative for emergence and positive for biomass. 5. Litter may have a positive effect on seedling recruitment in dry grasslands or under water-limited conditions, or in the presence of low to medium litter amounts (< 500 g m À2 ). However, high litter amounts (> 500 g m À2 ) will inhibit seedling recruitment. Large seeds showed a more positive response to litter presence with respect to seedling emergence and survival, but not concerning biomass. 6. Synthesis. Under dry conditions (e.g. dry grasslands or dry periods) or with low to medium litter amounts, litter presence has a positive effect on seedling establishment. However, climate and land use change may promote litter accumulation and reduce seedling establishment, affecting grasslands composition and ecosystem functions.

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Non-native plant species may contain allelopathic substances that might help to out-compete native vegetation. These allelochemicals may be released from live or dead plant tissues and be accumulated in the soil. We tested whether non-native species leaf litter and their leachates reduced seedling establishment and growth of native species. We subjected seeds of six native species to the effect of litter leachates of three of the most important invasive plants in Europe and to mannitol solutions with similar osmotic potential in germination chamber experiments. Additionally, we measured the effect of the same litter on emergence and growth of the native species in an outdoor pot experiment. Litter leachates delayed and reduced germination and affected initial root growth of all native species. The effects of leachates were significantly higher than those of mannitol, indicating the action of toxic, most probably allelochemical substances. Emergence of seedlings in pots was also reduced, but total biomass per pot was not affected and biomass per seedling increased. Allelochemicals may affect germination and early stages of seedling recruitment. However, these negative effects seem to cease shortly after germination, when other mechanisms such as competition may be more important. Consequently, litter-borne allelochemicals are unlikely to drive the invasion of the studied non-native species, but they may contribute to maintain mono-dominant stands reinforcing invasion success.

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co-occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.