Effects of litter amount and seed sowing position on seedling emergence and growth of hemiparasitic Rhinanthus species under drought stress.

Roadside vegetation in Central Europe is mostly species-poor and dominated by a few grass species. Hemiparasitic plant species, including Rhinanthus spp., might effectively restrict grass growth, thereby making space for light-dependent herb species. Despite the significance of abiotic site conditions for plant establishment in general, their effects on Rhinanthus establishment are less well known. We investigated combined effects of water availability, litter amount and seed position within litter on Rhinanthus seedling emergence and growth. Two parallel greenhouse experiments were conducted with R. angustifolius and R. minor. In these, we tested the impact of 200 or 400 g litter·m-2 with seeds sown beneath or on top of a litter layer under constantly humid or intermittently dry conditions on seedling emergence and biomass production of Rhinanthus. Presence of litter positively affected Rhinanthus seedling emergence when sown beneath the litter layer and reduced negative effects of water deficiency. Sowing beneath a litter layer increased seedling emergence by 157%, with similar effects at 200 and 400 g litter·m-2. Water level did not affect biomass production. Compared to R. minor, R. angustifolius had higher mean biomass, and its seedlings emerged earlier and in higher numbers. Our results indicate that Rhinanthus spp. react similarly to litter as non-hemiparasitic plant species from temperate grasslands. Litter presence positively influenced Rhinanthus seedling emergence and growth under intermittently dry conditions. Its hemiparasitic characteristics might reduce drought impacts on biomass production. To ensure seed contact with the soil surface, seeds should be sown when no litter is present, or mulching should occur post-sowing.

Effects of macrocyclic lactone anthelmintics on seed germination of temperate grassland species.

Macrocyclic lactone anthelmintics are widely used to control invertebrate pests in livestock, such as sheep. While anthelmintic effects on non-target animals, such as dung-dwelling insects, are well studied, effects on seed germination are largely unknown. Seeds can come into contact with anthelmintics either during passage through the gastro-intestinal tract of grazing animals or when anthelmintics are excreted with their dung into the environment, which may result in changed germination patterns. We used four commonly applied macrocyclic lactones to assess their effects on germination: moxidectin, ivermectin, abamectin and doramectin as pure substances; moxidectin and ivermectin also in formulated form. We tested these pharmaceuticals on 17 different temperate grassland species from five plant families. Seeds were exposed to three concentrations of macrocyclic lactones (0.1, 1.0 and 10.0 mg·l-1 ) under controlled conditions, and germination was assessed over a 6-week period. From these data, we calculated germination percentage, mean germination time and germination synchrony. Most of the tested species were significantly affected in germination percentage and/or mean germination time by at least one of the tested pharmaceuticals, with formulated moxidectin having the largest impact. In general, the effects found were species- and pharmaceutical-specific. While formulated substances generally reduced germination percentage and increased mean germination time, pure substances increased germination percentage. Synchrony showed less clear patterns in all pharmaceuticals. Although effect size and sign varied between species, our study shows that non-target effects of macrocyclic lactones commonly occur in terrestrial plants. This may impede successful seed exchange between habitats via sheep, and even translate into profound changes to grazed ecosystems.

Differential effect of drought regimes on the seedling performance of six floodplain grassland species.

The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root-mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands. In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results. In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species-specific (V. maritima being the most sensitive species) and habitat-specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands. Because of species- and habitat-specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

Distinct germination response of endangered and common arable weeds to reduced water potential.

Arable weeds are one of the most endangered species groups in Europe. Modern agriculture and intensive land-use management are the main causes of their dramatic decline. However, besides the changes in land use, climate change may further challenge the adaptability of arable weeds. Therefore, we investigated the response pattern of arable weeds to different water potential and temperature regimes during the phase of germination. We expected that endangered arable weeds would be more sensitive to differences in water availability and temperature than common arable weeds. To this end, we set up a climate chamber experiment where we exposed seeds of five familial pairs of common and endangered arable weed species to different temperatures (5/15, 10/20 °C) and water potentials (0.0 to -1.2 MPa). The results revealed a significant relationship between the reaction of arable weed species to water availability and their Red List status. The effects of reduced water availability on total germination, mean germination time and synchrony were significantly stronger in endangered than in common arable weeds. Therefore, global climate change may present a further threat to the survival of endangered arable weed species.

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species.

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.

Litter effects on seedling establishment interact with seed position and earthworm activity.

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 × 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.