Rhinanthus minor population genetic structure and subspecies: Potential seed sources of a keystone species in grassland restoration projects

In the last few decades unimproved semi-natural grasslands have been affected by intensification of land use and habitat fragmentation. Because of their biodiversity these species-rich grasslands are of high conservation importance and efforts are under way to restore such habitats. Detailed knowledge of within species diversity will aid deciding on the optimal seed source for such restoration projects, e.g. local genotypes or ecotypes. Rhinanthus minor is a species that is typically found in semi-natural grasslands and is commonly used in grassland restoration projects. This is because R. minor is a hemiparasitic plant that takes minerals and nutrients from its host, which in turn decreases the host's biomass and leads to opportunities for less competitive species in the vegetation. Here, we investigate genetic diversity within and between R. minor populations. This allowed us to test whether the six different subspecies of R. minor that have been described in the UK, based on their morphology, flowering time, and habitat, can be differentiated using molecular markers. We identified moderate levels of genetic differentiation between R. minor populations within the UK. In addition, R. minor individuals from the UK appear to be distinct from R. minor and Rhinanthus angustifolius individuals from other European countries based on microsatellite genotyping and DNA sequencing of cpDNA and rDNA ITS. The molecular markers used in the current study did not separate populations of R. minor based on either their subspecies or habitat. The implication for the use of R. minor in grassland restoration projects seems to be that it is not necessary to use local seeds or seeds from the same subspecies.     

Closely related parasitic plants have similar host requirements and related effects on hosts

The performance of root hemiparasites depends strongly on host species identity, but it remains unknown whether there exist general patterns in the quality of species as hosts for hemiparasites and in their sensitivity to parasitism. In a comparative approach, the model root hemiparasites Rhinanthus minor and R. alectorolophus were grown with 25 host species (grasses, forbs, and legumes) at two nutrient levels. Hosts grown without parasites served as a control. Host species identity strongly influenced parasite biomass and other traits, and both parasites grew better with legumes and grasses than with forbs. The biomass of R. alectorolophus was much higher than that of R. minor with all host plants and R. alectorolophus responded much more strongly to higher nutrient availability than R. minor. The performance of the two species of Rhinanthus with individual hosts was strongly correlated, and it was also correlated with that of R. alectorolophus and the related Odontites vulgaris in previous experiments with many of the same hosts, but only weakly with that of the less closely related Melampyrum arvense. The negative effect of R. minor on host biomass was less strong than that of R. alectorolophus, but stronger relative to its own biomass, suggesting that it is more parasitic. The impact of the two parasites on individual hosts did not depend on nutrient level and was correlated. Several legumes and grasses were tolerant of parasitism. While R. minor slightly reduced mean overall productivity, R. alectorolophus increased it with several species, indicating that the loss of host biomass was more than compensated by that of the parasite. The results show that closely related parasites have similar host requirements and correlated negative effects on individual hosts, but that there are also specific interactions between pairs of parasitic plants and their hosts.     

Interactions of the Hemiparasitic Species Rhinanthus minor with its Host Plant Community at Two Nutrient Levels

For root hemiparasites, host plants are both the source of water and nutrients below-ground, but competitors for light above-ground. Hemiparasites can reduce host biomass, and in this way considerably affect the whole plant community. To investigate these effects, we carried out two experiments in an oligotrophic meadow with a native population of Rhinanthus minor. In the first experiment, removal of R. minor was combined with fertilization in a factorial design, and in the second one, we manipulated R. minor density by thinning. The presence of R. minor decreased the biomass of its host community, mostly by suppressing grasses. In this way, the species was able to counterbalance the effect of fertilization, which increased community biomass and in particular that of grasses. Neither the presence of R. minor nor fertilization affected the total number of species or the Shannon-Wiener diversity index (H’) of the host community. However, H’ of grasses was higher and H’ of forbs (non-leguminous dicots) was lower in the presence of R. minor. Reduction of grasses by R. minor favored mainly the dominant forb Plantago lanceolata, which partly acquired the role of a competitive dominant. Effects of R. minor on community diversity seem to be highly dependent on the relative sensitivities of dominant and subordinate species. Fertilization increased the mortality of seedlings, resulting in a lower number of flowering plants. However, surviving individuals on average produced more flowers. Thinning resulted in lower mortality of R. minor plants. This indicates that intraspecific competition in R. minor populations results in negative density dependence.     

Hybridization in Annual Plants: Patterns and DynamicsDuring a Four-Year Study in Mixed Rhinanthus Populations

Hybridization in annual plants is rare, but their short life cycle provides an excellent opportunity to study the dynamics of hybridization. Hybridization occurs between the annual hemiparasites Rhinanthus minor and Rhinanthus angustifolius (Orobanchaceae). Using flower morphology, Kwak (1980) found a prevalence of hybrids close to R. angustifolius in a single population. We aim to find whether this pattern is also found using genetic markers, whether it is generally occurring in mixed populations, and whether these populations are stable over time. We used species-specific genetic markers to determine the number of individuals in a range of hybrid classes in three mixed populations of different ages during four consecutive years. In the young population, F₁ hybrids were found in the first year and mostly hybrids between R. minor and these F₁s in the second year, but in the years after that, hybrids close to R. angustifolius became more abundant. We also found this in the two older populations, where hybrids close to R. angustifolius always occurred in higher frequencies than hybrids close to R. minor. Over time, R. angustifolius strongly increased in frequency in two populations. Patterns of marker presence and absence suggested that advanced-generation hybrids are mainly formed by backcrossing with one of the parents, predominantly R. angustifolius whenever its frequency in the population is higher than 15%. The dynamics of mixed populations depend on the ecological conditions that regulate the presence of the two parental species, and introgression into R. angustifolius seems prevalent.     

Long‐term dynamics of the hemiparasite Rhinanthus angustifolius and its relationship with vegetation structure

Questions: 1. How are the long‐term dynamics of the root hemiparasite Rhinanthus angustifolius related to vegetation structure, grassland management and climate? 2. Does R. angustifolius have a long‐term impact on standing crop and community composition? Location : A formerly fertilized grassland, part of a larger brook valley system in the nature reserve ‘Drentsche Aa’, near Groningen, The Netherlands. Methods : Vegetation development has been monitored since 1973 in 54 permanent plots in nine management regimes without fertilizer application. Results : 1. The hemiparasite established when standing crop was less than 600 g.m^?2 and performed best under annual haymaking using machinery. Since its appearance, the population fluctuated stochastically, with two peaks. Coinciding collapses in six adjacent grasslands and comparison with an integrated climatic index suggest that the population collapses are induced by spring drought. 2. We did not find a relationship between total standing crop and R. angustifolius cover. Cover of grasses was negatively related to the abundance of the hemiparasite in the same and the previous year. Forb cover tended to increase with the parasite. Conclusions : R. angustifolius shows stochastic population fluctuations, mainly determined by spring drought, to which this species is probably highly vulnerable because of its parasitic and annual life style without a persistent seed bank. The hemiparasite also shows long‐term relationships with grass cover (negative) and forb cover (positive), but it seems to have no lasting impact on standing crop.     

Outcrossing rates in two self‐compatible,hybridising Rhinanthus species: implications for hybrid formation

The congeners Rhinanthus angustifolius and Rhinanthus minor, two annual hemiparasites pollinated by bumblebees, are known to hybridise in the wild. Both species are self‐compatible, but the capacity for autonomous selfing is higher in R. minor. This suggests a difference in realized outcrossing rates, which have not been determined before in these species. Using microsatellites, both species turned out to have mixed mating systems, but with a much lower multilocus outcrossing rate (0.13) for R. minor compared to R. angustifolius (0.76). We hypothesised that a higher outcrossing rate should lead to a higher chance of heterospecific pollination, and we therefore determined the rate of hybrid formation on each species in an artificial mixed population. Hybrid seeds were produced at low frequency (4.5%), and no significant difference was found between the species. It is therefore likely that post‐pollination processes influence hybrid seed formation to counteract the expected difference in heterospecific pollen deposition. We checked fruit set, seed set and the rate of autonomous selfing in controlled crosses in the greenhouse in 2 years, and found that fruit set (2003) or seed set (2010) were lower in R. angustifolius × R. minor crosses relative to the reciprocal cross. Hybrid seeds produced on R. angustifolius also had a much lower germination rate, so most of the established F₁ hybrid plants have the R. minor cytoplasm. The formation of advanced hybrids depends on pollinator preference, which is biased towards R. angustifolius if present in sufficient numbers, because it offers more rewards.     

Parasitic plants—impacts on nutrient cycling

Whilst it is widely accepted that different plant species influence ecosystem processes such as nutrient cycling, we are still far from understanding and predicting the effects of changes in vegetation composition on biogeochemical cycles. Ameloot E, Verlinden G, Boeckx P, Verheyen K, Hermy M. Impact of hemiparasitic Rhinanthus angustifolius and Rhinanthus minor on nitrogen availability in grassland (this issue) examine, for the first time, the impact of root hemiparasitic plants on in-situ nitrogen dynamics in intact field vegetation, using ¹⁵N techniques. This represents an important step forward in elucidating the unusual roles of parasitic plants in ecosystem function, and builds on a small set of recent studies indicating that parasitic plants have the potential to substantially alter the quantity and quality of resources returned to the soil by the plant community. These results reinforce the need to include parasitic plants in community and ecosystem theory, and emphasise that understanding the roles of parasitic plants in ecosystems is relevant for understanding ecosystem dynamics in a changing world.     

Effects of two contrasting hemiparasitic plant species on biomass production and nitrogen availability

Hemiparasitic plants can substantially change plant community structure; the drainage of host resources has a direct negative effect on host biomass and, as a consequence, promotes non-host biomass production (parasitism pathway); on the other hand, hemiparasitic litter inputs can enhance nutrient cycling which may have an indirect positive effect on both host and non-host biomass production (litter pathway). We evaluated the net effect of both pathways on total shoot biomass (with and without the hemiparasite) and shoot biomass of graminoids, forbs and ericaceous shrubs using a removal experiment in three sites infested with the annual Rhinanthus angustifolius, and three sites infested with the biennial Pedicularis sylvatica. We addressed the potential importance of litter effects by determination of litter quantity and quality, as well as modeling N release during decomposition. In the second year after removing the hemiparasites, total plant biomass at Rhinanthus sites was 24 % higher in weeded plots than in control plots, while weeding had no significant effect at Pedicularis sites. The increase in total biomass following Rhinanthus removal was mainly due to a higher biomass of graminoids. The amount of litter produced by Rhinanthus was only half of that produced by Pedicularis; N contents were similar. The amount of N in the litter was 9 and 30 % of the amount removed by mowing for Rhinanthus and Pedicularis sites, respectively. Within 2 months, about 45 % of the N in both hemiparasitic litter types was released by decomposition. Our results suggest that in addition to the suppression of host biomass due to parasitism, positive litter feedbacks on host and non-host biomass—via an increase in nutrient availability—also affect plant community structure. We propose that, depending on the particular hemiparasite and/or site conditions, these positive litter feedbacks on shoot biomass can compensate for the negative effect of parasitism.     

Shared pollinators and pollen transfer dynamics in two hybridizing species, Rhinanthus minor and R. angustifolius

Gene flow between hybridizing plant species depends strongly on pollinator behaviour, which affects pollen transfer among floral types and reproductive isolation. We examined bumblebee behaviour and pollen transfer between two hybridizing Rhinanthus species that are very similar in ecology and floral traits. The two species, Rhinanthus minor and R. angustifolius, shared similar pollinator guilds and assemblages, but pollinator recruitment and flower visitation rates were higher in R. angustifolius sites, probably because of its higher reward levels and better visibility. When presented with Rhinanthus flowers, bumblebees that previously foraged on R. angustifolius were less prone to visit R. minor inflorescences, while R. minor foragers accepted both species in similar proportions. Although Rhinanthus has been cited as a case of mechanical isolation resulting from interactions between bee behaviour and differences in stigma and anther placement, we found no support for efficient mechanical reproductive isolation. Bumblebees that foraged on R. minor flowers carried more pollen, but pollen placement on their bodies was similar to that of bees that visited R. angustifolius, and cross-specific stigmatic pollen deposition was similar in both directions. However, the asymmetry in pollinator handling time between the two species, due to dissimilar pollen rewards, may have lowered relative heterospecific pollen receipt on R. angustifolius, suggesting that net gene flow resulting from pollen transfer dynamics is more likely towards R. minor, although this effect remains weak and will be most likely counterbalanced by context-based labile pollinator preference.     

Effects of grazing intensity and grazing exclusion on litter decomposition in the temperate steppe of Nei Mongol,China

Aims Grazing intensity and grazing exclusion affect ecosystem carbon cycling by changing the plant community and soil micro-environment in grassland ecosystems. The aims of this study were: 1) to determine the effects of grazing intensity and grazing exclusion on litter decomposition in the temperate grasslands of Nei Mongol; 2) to compare the difference between above-ground and below-ground litter decomposition; 3) to identify the effects of precipitation on litter production and decomposition. Methods We measured litter production, quality, decomposition rates and soil nutrient contents during the growing season in 2011 and 2012 in four plots, i.e. light grazing, heavy grazing, light grazing exclusion and heavy grazing exclusion. Quadrate surveys and litter bags were used to measure litter production and decomposition rates. All data were analyzed with ANOVA and Pearson’s correlation procedures in SPSS. Important findings Litter production and decomposition rates differed greatly among four plots. During the two years of our study, above-ground litter production and decomposition in heavy-grazing plots were faster than those in light-grazing plots. In the dry year, below-ground litter production and decomposition in light-grazing plots were faster than those in heavy-grazing plots, which is opposite to the findings in the wet year. Short-term grazing exclusion could promote litter production, and the exclusion of light-grazing could increase litter decomposition and nutrient cycling. In contrast, heavy-grazing exclusion decreased litter decomposition. Thus, grazing exclusion is beneficial to the restoration of the light-grazing grasslands, and more human management measures are needed during the restoration of heavy-grazing grasslands. Precipitation increased litter production and decomposition, and below-ground litter was more vulnerable to the inter-annual change of precipitation than above-ground litter. Compared to the light-grazing grasslands, heavy-grazing grasslands had higher sensitivity to precipitation. The above-ground litter decomposition was strongly positively correlated with the litter N content (R² = 0.489, p < 0.01) and strongly negatively correlated with the soil total N content (R² = 0.450, p < 0.01), but it was not significantly correlated with C:N and lignin:N. Below-ground litter decomposition was negatively correlated with the litter C (R² = 0.263, p < 0.01), C:N (R² = 0.349, p < 0.01) and cellulose content (R² = 0.460, p < 0.01). Our results will provide a theoretical basis for ecosystem restoration and the research of carbon cycling.     

内蒙古温带草原不同放牧强度和围栏封育对凋落物分解的影响

放牧和围封通过影响植物群落结构和土壤微环境来调控草地生态系统的碳循环。该研究在内蒙古温带草原设置轻度放牧后围封、轻度放牧、重度放牧后围封、重度放牧4种样地, 通过测定干旱年(2011年)和湿润年(2012年)地上、地下凋落物产量、质量及其分解速率和土壤养分含量, 分析不同放牧强度对凋落物形成和分解的影响, 以及围栏封育对生态系统恢复的作用。结果表明: 重度放牧地上凋落物产量和分解速率均高于轻度放牧。干旱年轻度放牧样地地下凋落物产量和分解速率高于重度放牧, 湿润年相反。短期围封显著提高了凋落物产量, 轻度放牧样地围封后地上凋落物分解速率和养分循环加快, 而重度放牧样地围封后地上凋落物分解减慢。因此, 与重度放牧相比, 轻度放牧草地的恢复更适合采用围栏封育措施; 而重度放牧草地的恢复可能还需辅以必要的人工措施。降水显著促进地上、地下凋落物形成和分解。地下凋落物的生产和分解受降水年际波动影响较大, 重度放牧草地对降水变化的敏感度比轻度放牧草地高。地上凋落物分解速率与凋落物N含量显著正相关, 与土壤全N显著负相关, 与地上凋落物C:N和木质素:N相关性不大; 地下凋落物分解速率与凋落物C、C:N和纤维素含量显著负相关。该研究结果将为不同放牧强度的草地生态系统恢复和碳循环研究提供理论依据。     

Meta-analysis of standing crop reduction byRhinanthus spp. and its effect on vegetation structure

We performed a quantitative literature review on the effect of the root hemiparasiteRhinanthus on vegetation standing crop. (1) Across all available experimental studies in mixed vegetation and in pots, above-ground biomass of co-occurring species is generally reduced, with on average 40% and 60% of the value in the controls respectively. Total above-ground biomass, as the sum of parasite biomass and biomass of co-occurring species, decreases in most cases. For field experiments this reduction amounts, on average, to 26% of the control value. This implies that there is no compensation by the parasites’ biomass for the loss of biomass of co-occurring species due to parasite infection. This can be attributed to the low resource-use efficiency of hemiparasites. Meta-analysis confirmed these trends. (2) In pot experiments, the negative effect of the parasite on the above-ground biomass of the host increases with the number ofRhinanthus plants. In field experiments, we found no relationship between biomass reduction andRhinanthus density. (3) Total above-ground biomass reduction in field experiments increases with standing crop of the vegetation. However, reduction in above-ground biomass of co-occurring species seems to decrease with standing crop. Functional and species diversity buffer the community against negative effects ofRhinanthus. (4) In field experiments, functional groups are affected differently byRhinanthus spp. Grasses and legumes are mostly strongly reduced by the hemiparasites. Non-leguminous dicots mostly benefit from the presence ofRhinanthus. (5) In one out of four weeding experiments,Rhinanthus spp. has a significant (positive) effect on species number. However, the response of plant diversity to invasion of parasitic plants requires further research.     

Nutrient input from hemiparasitic litter favors plant species with a fast-growth strategy

Aims

Hemiparasitic plants often produce nutrient-rich litter with high decomposition rates, and thus can enhance nutrient availability. When plant species have differential affinities for this nutrient source, hemiparasitic litter might influence species composition in addition to the parasitic suppression of host species. We expected that species adapted to fertile habitats derive a higher proportion of nutrients from the hemiparasitic litter compared to other species.

Methods

¹⁵N-labeled litter of Rhinanthus angustifolius and Pedicularis sylvatica was added to experimental field plots and adjacent litter bags. We examined N release from the litter, N uptake by the vegetation 2, 4 and 12 months after litter addition and differences in the proportion of N taken up from the litter (N_L) between co-occurring species.

Results

The percentage of N in shoots of co-occurring plant species that is derived from the added hemiparasitic litter (N_L) strongly differed between the species (0.1–6.2 %). After exclusion of species with an alternative N source (legumes as well as ectomycorrhizal and ericoid mycorrhizal species), N_L was positively related (p?<?0.001) with specific leaf area (SLA) and at Pedicularis sites with leaf N concentration (LNC) and leaf phosphorus concentration (LPC) (p?<?0.05), i.e. leaf traits associated with a fast-growth strategy and adaptation to high-nutrient environments.

Conclusions

Our results suggest that nutrient release from hemiparasitic litter favors plant species with a fast-growth strategy adapted to high-nutrient environments compared to species with a slow-growth strategy. Whether continued hemiparasitic litter inputs are able to change species composition in the long term requires further research.     

Detecting hybridization in mixed populations ofRhinanthus minor andRhinanthus angustifolius

Rhinanthus minor andRhinanthus angustifolius are known to hybridize in mixed populations in nature. These hybridization events can have important evolutionary consequences. The development and use of species-specific RAPD and ISSR markers allowed the detection of hybrid individuals not always distinguishable with morphological characters. Two mixed populations of different ages were studied. In a young mixed 2-year-old population, both individuals of the two parental species and F₁ hybrids were found using genetic analysis, showing that hybridization occurred rapidly. Flower morphology of F₁ hybrids was too variable to distinguish all these hybrids from the parental species. This morphological variability of F₁ hybrids was also confirmed in artificial crosses in the greenhouse. In an old and no longer mixed 30-year-old population, onlyR. angustifolius plants and a few genetically introgressed individuals close toR. angustifolius were present. Genetic markers showed traces of past hybridization and introgression. Unidirectional introgression ofR. minor intoR. angustifolius with the complete disappearance ofR. minor from this population was observed.     

Invasion by <Emphasis Type="Italic">Aegilops triuncialis</Emphasis> (Barb Goatgrass) Slows Carbon and Nutrient Cycling in a Serpentine Grassland

Invasive plant species alter plant community composition and ecosystem function. In the United States, California native grasslands have been displaced almost completely by invasive annual grasses, with serpentine grasslands being one of the few remaining refugia for California grasslands. This study examined how the invasive annual grass, Aegilops triuncialis, has altered decomposition processes in a serpentine annual grassland. Our objectives were to (1) assess howA. triuncialis alters primary productivity and litter tissue chemistry, (2) determine whether A. triuncialis litter is more recalcitrant to decomposition than native litter, and (3) evaluate whether differences in the soil microbial community in A. triuncialis-invaded and native-dominated areas result in different decomposition rates of invasive and/or native plant litter. In invaded plant patches, A. triuncialis was approximately 50% of the total plant cover, in contrast to native plant patches in which A. triuncialis was not detected and native plants comprised over 90% of the total plant cover. End-of-season aboveground biomass was 2-fold higher in A. triuncialis dominated plots compared to native plots; however, there was no significant difference in belowground biomass. Both above- and below-ground plant litter from A. triuncialis plots had significantly higher lignin:N and C:N ratios and lower total N, P, and K than litter from native plant plots. Aboveground litter from native plots decomposed more rapidly than litter from A. triuncialis plots, although there was no difference in decomposition of belowground tissues. Soil microbial community composition associated with different soil patch types had no effect on decomposition rates. These data suggest that plant invasion impacts decomposition and nutrient cycling through changes in plant community tissue chemistry and biomass production.     

Contrasting effects of hemiparasites on ecosystem processes: can positive litter effects offset the negative effects of parasitism?

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castilleja occidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes.     

Opposite effects of litter and hemiparasites on a dominant grass under different water regimes and competition levels

Direct and indirect biotic interactions may affect plant growth and development, but the magnitude of these effects may vary depending on environmental conditions. In grassland ecosystems, competition is a strong structuring force. Nonetheless, if hemiparasitic plant species are introduced the competition intensity caused by the dominant species may be affected. However, the outcome of these interactions may change between wet or dry periods. In order to study this, we performed a pot experiment with different densities of the dominant species Schedonorus arundinaceus (1, 2 or 4 individuals) under constantly moist or intermittently dry conditions. The different Schenodorus densities were crossed with presence or absence of hemiparasites (either Rhinanthus minor or R. alectorolophus). Additionally, pots remained with bare ground or received a grass litter layer (400 g m^?2). We expected that indirect litter effects on vegetation (here Schedonorus or Rhinanthus) vary depending on soil moisture. We measured Schedonorus and Rhinanthus aboveground biomass and C stable isotope signature (δ¹³C) as response variables. Overall, Schedonorus attained similar biomass under moist conditions with Rhinanthus as in pots under dry conditions without Rhinanthus. Presence of Rhinanthus also increased δ¹³C in moist pots, indicating hemiparasite-induced water stress. Litter presence increased Schedonorus biomass and reduced δ¹³C, indicating improved water availability. Plants under dry conditions with litter showed similar biomass as under wet conditions without litter. Hemiparasites and litter had opposite effects: hemiparasites reduced Schedonorus biomass while litter presence facilitated grass growth. Contrary to our expectations, litter did not compensate Schedonorus biomass when Rhinanthus was present.     

Recovery of plant species composition and ecosystem function after cessation of grazing in a Mediterranean grassland

Short- and long-term changes in species composition, plant biomass production, and litter decomposition after cessation of grazing were examined in a Mediterranean grassland with high dominance of annual species and strong seasonality in biomass production. Short-term changes were assessed during three consecutive years in plots previously exposed to different grazing pressures and compared to plots in long-term (30–40 years) exclosures. Short-term cessation of grazing led in the short-term to an increase in relative biomass of annual crucifers and tall annual and perennial grasses, while biomass of annual legumes, annual thistles and short annual grasses decreased. Consequently, similarity increased between vegetation recently excluded from grazing and vegetation in long-term protected plots. Our research showed that in systems with high dominance of grasses and annual species, the rapid changes in plant species composition that occur after grazing cessation were associated with a fast recovery of the potential for biomass production to levels found in long-term protected plots, while litter decomposition rate did not change even after long-term cessation of grazing. Moreover, previous history of grazing did not affect plant litter decomposition, despite higher litter quality in grazed treatments. This study provides new insights about the processes involved in the diverse responses of ecosystem functions resulting from shifts in species composition associated with grazing cessation and land use change in Mediterranean grasslands.     

Seeds of change: The value of using Rhinanthus minor in grassland restoration

Question: What is the value of using Rhinanthus minor in grassland restoration and can restrictions on its establishment be overcome? Location: England (United Kingdom). Methods: Two experiments were established to determine the efficacy of inoculating R. minor on a suite of four agriculturally improved grasslands and the efficacy of using R. minor in grassland restoration. In Experiment 1, the effect of herbicide gap creation on the establishment and persistence of R. minor in grasslands ranging in productivity was investigated with respect to sward management. In Exp. 2, R. minor was sown at 1000 seeds/m² in conjunction with a standard meadow mix over a randomized plot design into Lolium perenne grassland of moderate productivity. The treatment of scarification was investigated as a treatment to promote R. minor. Results: Gap size had a significant role in the establishment and performance of R. minor, especially the 30 cm diameter gaps (Exp. 1). However, R. minor failed to establish long‐term persistent populations in all of the agriculturally improved grasslands. In Exp. 2, establishment of R. minor was increased by scarification and its presence was associated with a significant increase in Shannon diversity and the number of sown and unsown species. Values of grass above‐ground biomass were significantly lower in plots sown with R. minor, but values of total above‐ground biomass (including R. minor) and forb biomass (not including R. minor) were not affected. Conclusions: The value of introducing R. minor into species‐poor grassland to increase diversity has been demonstrated, but successful establishment was dependent on grassland type. The scope for using R. minor in grassland restoration schemes is therefore conditional, although establishment can be enhanced through disturbance such as sward scarification.     

Effect of nitrogen addition and drought on above-ground biomass of expanding tall grasses <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis> and <Emphasis Type="Italic">Arrhenatherum elatius</Emphasis>

Tall expansive grasses act as serious weeds since they spread intensively and are a important threat to biodiversity of various plant communities. A field experiment was set up based on three sets of paired plots, where Calamagrostis epigejos and Arrhenatherum elatius dominated and a mixture of both these tall grasses occurred. Parallel plots were treated by additional amounts of nitrogen (50 kg N ha⁻¹) for six years (2002–2007). Above-ground biomass was harvested in both wet (2004) and dry (2007) years, separated in individual fractions, and weighed to determine the above-ground biomass. We wanted to know if a higher nitrogen availability in dry year can support above-ground growth of expansive tall grasses C. epigejos and A. elatius and what their competitive ability of these tall grasses in mixture is in dry conditions. Drought resulted in a decrease in above-ground biomass production of studied tall expansive grasses, and, in a relative increase in mortality of A. elatius shoots in comparison with C. epigejos. Drought can substantially reduce the effect of applied nitrogen on above-ground growth of expansive tall grasses. However, A. elatius appears to be producing more above-ground biomass after application of N even in relatively dry conditions.