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Salinity is a potential environmental stress factor for plants. Improving plant growth under salinity stress requires an understanding of resistance mechanisms. Salinity stress resistance is related to both salinity avoidance and tolerance. In this study, interspecific differences in salinity stress resistance, avoidance, and tolerance were examined in eight C3 turfgrass species, namely Agrostis alba, Agrostis tenius, Dactylis glomerata, Festuca ovina, Festuca rubra, Lolium perenne, Phleum pratense, and Poa pratensis. In a greenhouse experiment, plants were exposed to 50, 100, or 200 mM NaCl for 2 weeks supplied via a hydroponics system. We found that interspecific differences in salinity stress resistance were associated mainly with salinity tolerance. Salinity avoidance mechanisms also contributed significantly to stress resistance. The contribution of genotype towards interspecific variation in salinity resistance and tolerance was higher than that of Na+ concentration. Salinity stress had negative effects on shoot dry mass, water status, and membrane stability. Relative water content was significantly correlated with salinity resistance, avoidance, and tolerance, whereas no significant correlation was observed for ion leakage. Mineral contents showed no clear contribution to salinity tolerance. Species varied in their proline content even in control conditions, and there was a positive association between proline content and salinity resistance and tolerance, but a negative association between proline content and salinity avoidance. F. ovina and P. pratense displayed the highest and lowest salinity stress resistance, respectively. The results of this study suggest that both salinity tolerance and avoidance mechanisms contribute towards salinity resistance, and that variation in salinity stress resistance among species is attributed to differences in proline content.

Shrub encroachment has taken place in many of China’s northern grasslands. This study attempts to answer the following questions: which plant communities are present in these shrub-encroached grasslands (SEGs)? What are the species richness and composition of these communities? Which environmental factors determine the spatial distribution thereof? We investigated the community characteristics of 255 SEG plots with a size of 20 m × 20 m at 69 locations across grasslands in northern China. In each plot, paired 1-m² quadrats were established within shrub patches and the neighbouring grassy matrix to record herb species composition. The quantitative characteristics (abundance, coverage and height) of herbs were measured in 0.25-m² subquadrats, and soil samples were collected in the quadrats. Two-way indicator species analysis (TWINSPAN), detrended canonical analysis (DCA) and canonical correspondence analysis (CCA) were used for community clustering and ordination. The DCA and TWINSPAN results suggested that the SEGs in northern China can be divided into six community types: Potentilla fruticosa + Carex atrofusca, Spiraea hypericifolia + Festuca ovina, Caragana acanthophylla + Stipa sareptana, Caragana microphylla + Leymus chinensis, Caragana microphylla + Stipa klemenzii and Caragana tibetica + Cleistogenes songorica. At a regional scale, climate and soil nutrients controlled the spatial patterns of species richness and community composition of the SEGs in northern China. Temperature exerted a negative impact, whereas precipitation and nutrients had positive effects on species richness. Among the environmental factors used, climate was the major controller of the variations in community structure. These results provide new insights into the community composition of SEGs in China and enrich the global dataset of SEGs.

The two factors defining male reproductive success in plants are pollen quantity and quality, but our knowledge about the importance of pollen quality is limited due to methodological constraints. Pollen quality in terms of chemical composition may be either genetically fixed for high performance independent of environmental conditions, or it may be plastic to maximize reproductive output under different environmental conditions. In this study, we validated a new approach for studying the role of chemical composition of pollen in adaptation to local climate. The approach is based on high‐throughput Fourier infrared (FTIR) characterization and biochemical interpretation of pollen chemical composition in response to environmental conditions. The study covered three grass species, Poa alpina, Anthoxanthum odoratum, and Festuca ovina. For each species, plants were grown from seeds of three populations with wide geographic and climate variation. Each individual plant was divided into four genetically identical clones which were grown in different controlled environments (high and low levels of temperature and nutrients). In total, 389 samples were measured using a high‐throughput FTIR spectrometer. The biochemical fingerprints of pollen were species and population specific, and plastic in response to different environmental conditions. The response was most pronounced for temperature, influencing the levels of proteins, lipids, and carbohydrates in pollen of all species. Furthermore, there is considerable variation in plasticity of the chemical composition of pollen among species and populations. The use of high‐throughput FTIR spectroscopy provides fast, cheap, and simple assessment of the chemical composition of pollen. In combination with controlled‐condition growth experiments and multivariate analyses, FTIR spectroscopy opens up for studies of the adaptive role of pollen that until now has been difficult with available methodology. The approach can easily be extended to other species and environmental conditions and has the potential to significantly increase our understanding of plant male function.

Mulching has increasingly become a standard method used in maintaining low-input lawns. This can be attributed to cost-savings, as these maintenance systems do not require loading and hauling of grass clippings. It is also possible to reduce the amount of nitrogen (N) fertilization and thus save on fertilizer costs. The effects of mowing management and rate of nitrogen fertilization on quality and growth of urban lawns were studied in a field trial during 2007–2012 on lawns newly sown in autumn 2006 and consisting of a grass–legume mixture ( Lolium perenne 25%, Poa pratensis 25%, Festuca rubra 30%, Festuca ovina 5%, Anthoxanthum odoratum 5%, Cynosurus cristatus 5%, Trifolium repens 3%, Lotus corniculatus 2%). Nitrogen was used at three application levels: 0, 50 and 100 kg of N ha −1 year −1 . Swards were cut to a height of 40 mm five times annually and mulched to the same height. Mulching significantly affected the botanical composition of lawns, wherein a decrease in the portion of legumes (from 32% to 19.1%) was observed. The returned grass matter supported a significant increase in the grass component in the lawns (from 57.8% to 69.7% of total). When the clippings were returned, quality remained the same. Returned clippings had a positive impact on colour and increased the height of cut swards significantly. Application of nitrogen decreased the share of legumes, supported the grass component, statistically increased quality, improved colour, and increased sward height. The results show that returning grass clippings through mulching is beneficial. In any case, it is necessary to apply at least a basal amount of mineral fertilizer in order to preserve the quality of urban lawns over the long term.

International audience; Plant–plant chemical interactions in forests can have a strong impact on the biodiversity and dynamics of these ecosystems, particularly in Mediterranean forests where plants exhibit a high secondary metabolite diversity. Allelopathic interactions in Mediterranean ecosystems have been mostly studied in the first stages of ecosystem dynamics, shrublands and pine forests, but little is known about these interactions in mature oak forests. In this study, the allelopathic effect of three main woody species of downy oak forests (Quercus pubescens, Acer monspessu-lanum and Cotinus coggygria) on germination and growth of two herbaceous species (Festuca ovina and Linum per-enne) was tested through aqueous extracts obtained from different leaf phenological stages (green, senescent and litter). The germination velocity of the two target species was inhibited by the aqueous extracts of senescent leaves from all the woody species. The growth of F. ovina seedlings was affected by aqueous extracts of green leaves of all the woody species, while the growth of L. perenne was only affected by aqueous extracts of green leaves of A. monspessulanum. This shows that (i) allelochemicals released by leaf leachates of the dominant woody species could control the dynamic of the herbaceous species, and then their potential competition with trees and (ii) allelo-pathic effects of woody species are related to their phenological stage and seem consistent with the development stage of target species.

Questions (1) When alpine vegetation is actively restored by seeding, how is vegetation cover influenced by seeding treatments and soil conditions? (2) How does the cover of species differ when they are seeded in a mixture and how is their response influenced by soil conditions? (3) Do individual species perform better or worse in a mixture than when sown separately? Location Hjerkinn, Dovrefjell, Norway. Methods In a factorial, randomized field experiment, we recorded the percentage cover of Festuca ovina, Luzula multiflora ssp. frigida, and Poa alpina seeded in four different soil types for three years after seeding. We seeded the three species separately and in a mixture in organic topsoil, peat soil, mineral fine soil and mineral coarse soil. We also recorded seedling emergence in a greenhouse experiment, using the same seeding treatments. Results In the field experiment, vegetation cover established fastest when F. ovina was sown in monoculture followed by the seed mixture. After three years, mean cover of F. ovina was 1.4 higher than mean P. alpina cover and more than three times higher than mean L. multiflora cover for single-species treatments, and four (P. alpina) and 15 (L. multiflora) times higher when the species were seeded together. L. multiflora germinated slowly in the greenhouse experiment, which could partly explain its poor field performance. In the field experiment, establishment was faster in organic soils than mineral soils for all seeding treatments. The largest difference between F. ovina and L. multiflora performance in the mixture treatment was found in the organic soil types, where overall cover was larger than in the mineral soils. In the organic soils, F. ovina was slightly facilitated in the mixture treatment, while the opposite was found for L. multiflora. Conclusion When the restoration goal is to quickly establish a vegetation cover, seeding monocultures of rapidly establishing species may be more effective than seeding mixtures even in alpine sites where interspecific facilitation may prevail. This article is protected by copyright. All rights reserved.

Establishing perennial plant communities on roadsides is a proactive approach to sustainable road management. The environmental and economic benefits of increasing desirable roadside vegetation include improving slope stabilization, soil conservation, and roadway safety while reducing erosion, maintenance costs, and invasive plants. This study evaluated the success of roadside revegetation on 16 sites in Idaho and on one site in adjacent southwest Montana. Study sites represent a diversity of climatic, topographic, and edaphic conditions in order to provide a variety of examples of roadside revegetation projects in 6 different ecoregions of Idaho. We measured all species’ canopy cover, including noxious weeds and other invasive plants. Of the seeded grass species, 21 of 27 established. Bluebunch wheatgrass ( Pseudoroegneria spicata (Pursh)A. Love [Poaceae]) and Idaho fescue ( Festuca idahoensis Elmer [Poaceae]) were the best-performing native grasses. Seeded grass species that consistently established but had a low mean canopy cover (2–5%) included sheep fescue ( Festuca ovina L.), western wheatgrass ( Pascopyrum smithii (Rydb.) A. Love [Poaceae]), and basin wildrye ( Leymus cinereus (Scribn. & Merr.) A. Love [Poaceae]). Seeded forbs and shrub species had low establishment success and low mean percent canopy cover. For example, no seeded forb species had more than 1% mean canopy cover and greater than 50% establishment when present in the seed mix. Seed mixes of 10 or fewer species resulted in ≥ 50% of the species establishing. Idaho noxious weeds were present in less than half of the sites (47%). Cheatgrass ( Bromus tectorum L. [Poaceae]), an invasive annual grass, occurred in 82% of the sites. These results provide species selection recommendations based on ecoregion for use in roadside revegetation projects in Idaho and portions of western Montana.

Symbiotic Epichloë species are fungal endophytes of cool-season grasses that can produce alkaloids with toxicity to vertebrates and/or invertebrates. Monitoring infections and presence of alkaloids in grasses infected with Epichloë species can provide an estimate of possible intoxication risks for livestock. We sampled 3,046 individuals of 13 different grass species in three regions on 150 study sites in Germany. We determined infection rates and used PCR to identify Epichloë species diversity based on the presence of different alkaloid biosynthesis genes, then confirmed the possible chemotypes with high-performance liquid chromatography (HPLC)/ultraperformance liquid chromatography-tandem mass spectrometry (UPLC-MS/MS) and gas chromatography-mass spectrometry (GC-MS) measurements. Infections of Epichloë spp. were found in Festuca pratensis Huds. (81%), Festuca ovina L. aggregate (agg.) (73%), Lolium perenne L. (15%), Festuca rubra L. (15%) and Dactylis glomerata L. (8%). The other eight grass species did not appear to be infected. For the majority of Epichloë-infected L. perenne samples (98%), the alkaloids lolitrem B and peramine were present, but ergovaline was not detected, which was consistent with the genetic evaluation, as dmaW, the gene encoding the first step of the ergot alkaloid biosynthesis pathway, was absent. Epichloë uncinata in F. pratensis produced anti-insect loline compounds. The Epichloë spp. observed in the F. ovina agg. samples showed the greatest level of diversity, and different intermediates of the indole-diterpene pathway could be detected. Epichloë infection rates alone are insufficient to estimate intoxication risks for livestock, as other factors, like the ability of the endophyte to produce the alkaloids, also need to be assessed. IMPORTANCE Severe problems of livestock intoxication from Epichloë-infected forage grasses have been reported from New Zealand, Australia, and the United States, but much less frequently from Europe, and particularly not from Germany. Nevertheless, it is important to monitor infection rates and alkaloids of grasses with Epichloë fungi to estimate possible intoxication risks. Most studies focus on agricultural grass species like Lolium perenne and Festuca arundinacea, but other cool-season grass species can also be infected. We show that in Germany, infection rates and alkaloids differ between grass species and that some of the alkaloids can be toxic to livestock. Changes in grassland management due to changing climate, especially with a shift toward grasslands dominated with Epichloë-infected species such as Lolium perenne, may result in greater numbers of intoxicated livestock in the near future. We therefore suggest regular monitoring of grass species for infections and alkaloids and call for maintaining heterogenous grasslands for livestock.

This study examined the effects of soil heavy metals, macronutrients, texture and pH as well as plant species richness and composition on soil respiration, enzymatic activity, microbial biomass, metabolic quotient (qCO2) and arbuscular mycorrhizal fungi (AMF) at sites of historical Zn–Pb mining. The study was conducted both on a large scale (65 heaps scattered over the area of 750 km2) and on a small scale (25 plots along two 48 m transects extending from heaps to adjacent fallow fields). Total concentrations of metals exceeded 400 (Cd), 20,000 (Pb) and 80,000 (Zn) mg kg−1 at the most polluted sites. Although they decreased along the heap-fallow direction, they still remained above environmental standards in fallow soils. In contrast, some soluble metal forms increased with the increasing distance from heaps. Soil organic matter had the strongest positive effect on most microbial parameters. Total and/or available heavy metals exhibited significant negative effects on microbial biomass, enzymatic activity and AMF, and a positive effect on qCO2. Organic matter alleviated negative effects of heavy metals on microorganisms; they were not observed where the increase in the contamination was accompanied by the increase in organic matter content. Plant species richness affected positively enzymatic activity and mycorrhization level. Plant species composition possibly contributed to the formation of soil microbial communities, but its effect was entangled in that of heavy metals as plant communities changed along pollution gradients (from metal-tolerant grasslands dominated by Festuca ovina to calcareous grasslands and ruderal communities at less polluted sites).

Shooting activities is an important source of Pb in contaminated soils. Lead accumulates in superficial soil horizons because of its low mobility, favouring its uptake by plants and representing a high transference risk to the trophic chain. A combination of phytoremediation with nanoremediation techniques can be used to recover firing range soils and decrease the mobility, bioavailability and toxicity of Pb. This study examines in depth the changes in Pb behaviour in firing range soils by adding hydroxyapatite nanoparticles (HANPs). These nanoparticles (NPs) may immobilise Pb and improve the quality of these areas. The use of HANPs and the Pb effects were assessed in three different species (Sinapis alba L., Lactuca sativa L. and Festuca ovina L.), focusing on their germination and early growth, through phytotoxicity assays. Single extractions with CaCl2 (0.01 M) in soils treated with HANPs show that these NPs retained Pb and reduced highly its availability and mobility. HR-TEM and TOF-SIMS were used to determine the interactions between HANPs and Pb, as well as with soil components. According to TOF-SIMS and HR-TEM/EDS analysis, Pb was mainly retained by HANPs but also associated lightly to organic matter, Fe compounds and silicates. Phytotoxicity assays exposed that S. alba, L. sativa and F. ovina were able to germinate and develop in the firing range soils despite the high available Pb contents before adding HANPs. After adding HANPs, Pb retention increased, favouring the germination and the growth of roots in the three species. These results suggest that HANPs can be used to decrease the availability and the toxicity of Pb without negative effects in the species growth. Accordingly, the combination of phytoremediation and nanoremediation techniques can be a great tool to stabilise these soils, avoiding the Pb transfer to nearby areas and its entry in the trophic chain.

Phosphogypsum (PG) is a by-product of phosphate fertilizer production. This study is the first to investigate Bromis inermis and Festuca ovina root morphology and colonization with arbuscular mycorrhizal fungi and dark septate endophytes (DSE) in pure PG and PG with a soil cap (soil). Root length density was significantly lower for plants on PG than on soil and both plant species were colonized with AMF and DSE, with lower frequency and intensity in PG. Phosphorus (20 times), calcium (35 times) and sulphur (50 times) were significantly higher in PG than in soil; resulting higher root tissue concentrations were significantly negatively correlated with mycorrhizal colonization for Festuca but not Bromus. Higher fungal colonization of Bromus, especially with DSE known to be prevalent in extreme environments, may be responsible for its better performance on PG relative to Festuca.

International audience; In recent years, plantations of fast-growing tree species have emerged as a possible way to meet the increasing demand for biomass for renewable energy in Europe. Agroforestry plantations including fast-growing tree species could be an attractive option because they reduce land competition for biomass and food production while providing forest benefits. Today, the species interactions that determine whether a given mixture will be more productive than the corresponding monocultures are still poorly understood. Our objective was to assess the performance of fast-growing trees, i.e. poplar (Populus nigra × P. deltoides) and alder (Alnus glutinosa), in association with herbaceous species either N 2-fixing species (succession alfalfa (Medicago sativa) and clover (Trifolium pratense)) or graminoids (succession of wheat (Triticum aestivum)-triticale (Triticosecale)-temporary grassland consisting of mixtures of perennial ryegrass (Lolium perenne) and fescue (Festuca ovina)), respectively, and in association together. We compared tree growth in mixed stands to growth in the respective tree monocultures. An experimental plantation composed of three blocks including the different treatments was set up in 2014 in northeastern France. In the forest mixture, poplar and alder were planted in alternating rows, and in the agro-forestry plots, every second line of trees was replaced by the crops. No fertilizers (of any kind) were used during the experiments. During six growing seasons, tree growth (height and diameter at breast height) was monitored monthly in the agroforestry stands, the forest mixture stand and the two tree monocultures. By the end of 2019, poplar stem height in association with N 2-fixing crops (alfalfa, then clover) was higher than in the forest mixture and the monoculture. During the warmest growing season with a marked dry period in June-July (2018), poplar height growth rate in the agroforestry treatment was between three and five times higher than for the poplars in the monoculture and the forest mixture. A facilitation process in the agroforestry plantations could explain this result due to a significant enrichment of the soil in nitrogen by the N 2-fixing crops. This result only appeared several years after tree planting, and management actions were necessary in order to control the strong competition between crops and trees during the early stages. In the forest mixture, poplars associated with alders were smaller than in their monoculture, in spite of a stratification of the canopies of the two species. For alders, tree growth did not appear to be affected by either an association with graminoids or with poplar.

In the Mediterranean region, endozoochorous seed dispersal is a potential alternative for the recovery of degraded or abandoned grasslands and shrublands. The objectives of this study were to quantify the number of seeds that pass through the goat gut and to determine whether this process affects seed germination in six common herbaceous species of the Mediterranean area ( Ornithopus compressus L., Biserrula pelecinus L., Medicago polymorpha L., Melilotus officinalis L. Pall, Festuca ovina L. and Dactylis glomerata L). We also examined whether the inclusion of the seeds in dung affected seed germination and seedling establishment and growth. The highest seed recovery rate was obtained 24–72 h after ingestion. The amount of retrieved seeds varied among species and was generally low, with significant differences among some leguminous and grass species. With the exception of M. officinalis , the quantity of retrieved legume seeds (1.5–6.8%) was greater than that of the grass seeds (0.9–1.1%). A significant relationship was found between recovery of the seeds and their hardness and volume. In all species, the recovered seeds presented significantly lower germinability than seeds that had not been consumed. In general, all recovered legume seeds showed higher average germination rate values (18–50%) compared to recovered grass seeds (0–14%). Seedling emergence was lower in intact than in crumbled dung pellets and, in most cases, the values of both of these were lower than that of unconsumed (control) seeds. No seedling emergence was recorded for seeds of B. pelecinus and O. compressus contained in intact or crumbled faeces at all sampling times. While the percentages of recovery and subsequent emergence found in this study are low in absolute terms, we can conclude that a significant number of seeds may become established from the dung and could be a potential alternative for the recovery of degraded or abandoned grazing areas.

Soils formed in metallic mines and serpentinite quarries, among other unfavourable features, have high levels of heavy metals. They can release into the environment causing surface and subsurface water contamination, uptake by plants, their accumulation in the food chain and adverse effects on living organisms. In this work, we studied the magnitude of the soils’ toxic effects not only on spontaneous plants but also on two species with phytoremediation potential. Several soils from two different exploitations were selected: a lead and zinc mine and a serpentinite quarry. Soils were characterized, and the pseudo-total and extractable contents of Co, Cr and Ni in soils from a serpentinite quarry were determined. The Cd, Pb and Zn pseudo-total and extractable contents were determined in soils developed in the Pb/Zn abandoned mine. Using a biotest, the chronic toxicity of the soil samples on higher plants was determined. Festuca ovina L., Cytisus scoparius (L.) Link., Sinapis alba L. and Brassica juncea L. were selected, the first two because they are spontaneous plants in the study areas and the last two because they have heavy metal phytoremediation potential. Pseudo-total contents of Co, Cr and Ni in the serpentinite quarry soils and of Zn, Pb and Cd in the Zn/Pb mine soils exceed generic reference levels. CaCl2 is the reactant that extracts the highest proportion of Co, Cr and Ni in the quarry soils and EDTA the largest proportion of Pb Zn and Cd content in the mine soils. The germination index values based on seed germination and root elongation bioassays revealed increasing plant sensitivity to the mine soils in the following order: B. juncea

14 structures of polycyclic aromatic hydrocarbons (PAHs) were identified in the organogenic soil horizon and plants of the south shrub tundra belt, at a background plot and at a coal mine. The polyarenes in soils and plants were mainly represented by light structures. On the background and polluted areas, mosses of Pleurozium schreberi, Polytrichum commune and grasses of Deschampsia caespitosa, Festuca ovina contained the highest PAH amounts. PAH bioaccumulation in plants is mainly dependent on their physiological features. 4 plant groups with different PAH bioaccumulation capabilities were identified on the basis of the calculated biological absorption coefficient. The toxicological activity of polyarenes for both soils and plants at the background and polluted areas was determined by heavy PAHs. The highest values of toxicological activity were observed for mosses and grassy plants whilst the lowest values were for shrubs. PAHs at the plots under study are mainly of petrogenic origin.

Roadside turfgrass mixtures are subject to a wide variety of extreme stresses including heat, drought, low nutrient availability, ice cover, and road salt exposure. The currently specified turfgrass seed mixtures for roadsides in Minnesota have not been based on the results of recent, designed experiments. A quantitative approach to evaluating turfgrass species mixtures for roadsides in Minnesota is needed in order to improve the current mixture recommendation. The objectives of this experiment were to (1) assess the performance of several mixtures of cool-season turfgrass for survival on roadsides in central Minnesota, (2) quantitatively evaluate the influence of individual species on the survival of turfgrass mixtures on roadsides, and (3) identify a suitable mixture of cool-season turfgrass species for roadsides in central Minnesota. In fall 2011, three replications of 51 cool-season turfgrass mixtures, comprising nine species of turfgrass, were established at two roadside locations in the metropolitan area surrounding St. Paul, MN, USA. Survival of the established mixtures was assessed using digital image analysis to determine percent living ground cover during both spring and summer 2012, and again using a grid-intersect method during spring 2013. Several mixtures were identified that performed significantly better than the current specified mix during spring or summer 2012. The log-odds of a plot retaining at least 60% living turfgrass cover in spring 2013 decreased by 1.78 for mixtures including tall fescue ( Festuca arundinacea L.). Hard fescue [ Festuca trachyphylla (Hack.) Krajina] and sheep fescue ( Festuca ovina L.) increased the log-odds of success by 0.95 and 0.96, respectively, and both estimates were significant at the 90% confidence level. Quantitative analysis of survival percentage in spring 2013 indicated that the best mixture for roadsides in central Minnesota comprises 20% slender creeping red fescue, 40% hard fescue, and 40% sheep fescue. These results can be used by public works officials to implement and improve roadside turfgrass mixtures in Minnesota.

The present study was conducted to increase phytoextraction efficiency of Festuca ovina L. in lead contaminated soil in the EDTA-assisted (0, 1.5, 3, 1.5+1.5, 3+3, 6 mmol kg), assess the best time of plant harvesting to increase Pb uptake and method of EDTA application to reduce Pb leaching risk. The results revealed that the greatest Pb uptake was observed in 3EDTA treatment. Therefore, 3mmolkg was used in the second step for assessing harvest time for 15, 30 and 45 days. Results showed that the concentration of Pb in plant tissues was increased with the passage of time and the best harvest time to achieve maximum removal of Pb was 60 days of the first harvest. In the third step to reduce leaching of Pb-chelate, 3mmolkg EDTA in five ways of single, double, triple, quadruplet, quintuplet were added to the soil. The results indicated that under quintuplet application, Pb content reached its minimum concentration in the soil and in the plant organs, the Pb concentration was maximum and metal concentration in the plant organs did not vary significantly when triple, quadruplet and quintuplet dosages were added (p

Butterflies belong to one of the most intensively studied invertebrate groups. However, detailed information on the larval ecology is still lacking in many species. This is especially true for the grass-feeding subfamily Satyrinae, to which our study species, the Woodland Ringlet (Erebia medusa Fabricius, 1787), belongs. At the habitat level, E. medusa exhibits a clear preference for abandoned grasslands with litter accumulation. In this study, we assessed the host-plant and oviposition preferences of E. medusa at the microhabitat level, i.e., within occupied habitat patches, in the Diemel Valley (Central Germany). Only two species of grass, Festuca ovina agg. and F. rubra agg., were systematically used for oviposition. Egg density was positively correlated with patchiness of herb layer, potential daily length of sunshine, and abandonment. However, the explanatory power of these parameters was low. We attribute this primarily to an ample supply of host plants growing under suitable conditions in the occupied habitat patches of E. medusa. The positive effect of abandonment is probably due to the fact that the average height of oviposition sites of E. medusa is greater than in most other satyrine species, which renders the eggs more vulnerable to mechanical disturbance. Moreover, thick layers of litter, which develop after the cessation of grazing or mowing, might be important as they modify the microclimate at the locations of the immature stages.

Native grasses are important for the enhancement of landscape. Agropyron cristatum, A.intermedium, Festuca ovina, Festuca arundinaceae, Cynodon dactylon, Bromus inermis and B. confinis are the potential sources of low maintenance turfgrasses for semi-arid regions. This greenhouse study examined the interaction of four different levels of soil water contents with 75, 50, and 25% field capacity (FC) and non-irrigation on physiological and biochemical responses of each accession. Total carbohydrate and sucrose contents of F. arundinaceae were increased significantly under non-irrigated treatment but decreased in the other genotypes. Proline, hydrogen peroxide, and total ascorbate increased in all the species under drought. Also, with decreased soil water content, H2O2 content increased up to 50% in F. arundinaceae. Protein content of F. arundinacea increased significantly under drought stress compared to the control; however, it decreased in other genotypes. Ascorbate peroxidase activities were increased in Festuca sp. and decreased in A. cristatum. It was found that F. arundinacea was able to maintain higher relative water content level and osmotic potential in non-irrigated drought treatment than B. inermis and A. cristatum. It may be speculated that the observed drought stress tolerance was associated with the ability of accumulating compatible solutes and H2O2 signaling cascade. Also, the high activity of ascorbate peroxidase resulted in protection against oxidative damage. It seems that this mechanism worked better in F. arundinacea to be used as low maintenance turfgrass.

11 species of grasses and 10 species of leguminous plants were tested for salt tolerance. The biotest of germination viability and capacity in soline water was performed. The results proved usefulness of using seeds as bioin-dicators and allowed to show the significant interspecific differences. The following species – Lolium perenne, L. multiflorum, Festuca pratensis, Arrhenetherum elatius, Trifolium resupinatum, T. incarnatum and Melilotus albus show high level of tolerance. The less tolerant species were: Festuca ovina, Agrostis alba, Trifolium repens, Lotus corniculatus, Medicago lupulina and Medicago sativa.

Aims From the light-competition hypothesis, competition for light is asymmetric and the observed increases in plant-size variability with increasingly denser canopies are primarily due to competition for light. Greater plant height provides pre-emptive access to light and produces increased height differences among species. The question is what produces these differences in plant height or height growth response among species in response to fertilization. Methods In 2009, a field experiment of N, P and N + P enrichments at three levels each was initiated in an alpine meadow on the northeast Qinghai- Tibet Plateau. Effects of fertilization on species richness, aboveground net primary production (ANPP), relative light intensity and plant height of different plant functional groups were determined. Festuca ovina (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb) and Geranium pylzowianum (upright forb) were selected as exemplars of each of the indicated functional groups. The N: C ratios in aboveground biomass, gibberellic acid (GA3) concentrations in leaves, plant heights and height relative growth rate (RGR) of these exemplar species were analyzed in detail. Important Findings Species richness of grasses significantly increased with increasing N + P levels. Species richness of legumes and upright forbs decreased after N and N + P additions. P addition had no significant effect on species richness. The effects of N + P addition on species richness and ANPP were consistently stronger than those of the single N or P fertilization. Reductions in species richness caused by nutrient addition paralleled the increases in ANPP and decreases in light intensity under the canopies, indicating indirect effect of nutrient addition on species richness via ANPP-induced light competition. The exemplar species that responded most positively to fertilization in height and RGR also displayed stronger increases in their GA3 content and N:C ratios. GA3 concentrations and N: C ratios were positively correlated with height RGR when the data were pooled for all species. The tallest and the fastest-growing grass, F. ovina, had the largest increase in N: C ratios and the highest leaf GA3 concentrations after nutrient addition. These results indicated that differential responses of GA3 concentrations and N: C ratios to fertilization were related to the inequality in plant heights among species.

Since the contribution of total belowground bud bank and different bud types to community regeneration has rarely been explored, the vegetative offspring recruited from different belowground bud types was investigated in four plant communities along a grassland degradation gradient in northeastern China (Inner Mongolia). This gradient, between 1000 and 1500 m a.s.l., has been caused by overgrazing. It is a Leymus chinensis steppe which occupies about 3.0×105 ha. Recruitment from tiller buds was dominant (>80%) in determining the total vegetative offspring density along the whole grassland degradation gradient. However, the proportional contribution of tiller-ramets to total ramet recruitment was significantly greater (P

Spoil heaps are the visible footprint of hydropower production, particularly in vulnerable alpine environments. Speeding up vegetation development by seeding commercial grass species has been a common restoration practice for the last 50 years, but we lack information on whether seeded species decline and allow native plant cover to develop. We visually estimated cover of native vascular plants and five seeded grass species (Agrostis capillaris, Festuca ovina, Festuca rubra, Schedonorus pratensis and Phleum pratense) on eight spoil heaps at different elevations (boreal-alpine zone) in western Norway. Spoil heap vegetation was censused twice (9-20 and 24-36 years after spoil heap construction); the undisturbed surrounding vegetation was also censused on the second occasion. Total cover on the spoil heaps showed some increase, but remained far below that in surrounding areas. Cover of seeded grass species in the surroundings was low (but not negligible), indicating suboptimal establishment ability. Seeded species usually covered less than 20 % of the spoil heaps, and only F. rubra, F. ovina and A. capillaris contributed substantially. Proportional cover indicated better initial establishment by seeded species, but their cover decreased between the censuses on all but the highest located spoil heap. The persistence of seeded grass species is problematic, and despite the decrease in proportional cover, they are likely to persist for decades on spoil heaps, posing a risk of invasion of surrounding areas. We therefore recommend replacing the practice of seeding with more appropriate restoration measures.

Tolerance of four cool-season turfgrass species sheep fescue (Festuca ovina), perennial ryegrass (Lolium perenne), colonial bentgrass (Agrostis tenuis) and rough bluegrass (Poa trivialis) to SO2 was studied by investigating their injury and physiological responses under SO2 stress for 14 days. Results showed that sheep fescue and perennial ryegrass had better SO2 tolerance than rough bluegrass and colonial bentgrass. Chlorophyll a and carotenoid contents were significantly higher in the SO2-tolerant turfgrass species (sheep fescue and perennial ryegrass) than in the SO2-sensitive turfgrass species (rough bluegrass and colonial bentgrass). Relatively lower levels of reactive oxygen species (ROS), malondialdehyde, and electrolyte leakage (EL) were observed in SO2-tolerant turfgrass species. Gradually increased antioxidant enzyme activities and proline content in SO2-tolerant turfgrass species could counteract such damages and harmful effects caused by ROS and EL, respectively. Sulfite reductase played an im...

BACKGROUND AND AIMS: There is no knowledge of winter plant phenology and its controlling factors on the Qinghai–Tibetan Plateau (QTP). Thus, we conducted a 4 year winter phenology and growth dynamics study in the alpine meadow on the eastern QTP. METHODS: From November 2013 to March 2017, the phenology of the ‘winter-growth’ and ‘winter-green’ species was recorded every 5 d. In November–February from 2014 to 2015, the above-ground biomass (AGB) in random plots was calculated to distinguish different growth patterns among winter growing species. The percentage of winter abundance relative to the summer population for forbs and the percentage of absolute coverage for grasses (W/S) were calculated to describe the importance of the winter population to the summer population. The soil moisture (SM) and soil temperature (ST) were used to explore the controlling factors on the AGB. Pearson’s correlation analysis between winter phenology data and environmental variables, including air temperature (T(air)), snow cover fraction (SCF), SM and ST, was used to investigate the factors affecting winter phenology during November–February from 2014 to 2017. KEY RESULTS: There were 107 species in total in the sites, including ten ‘winter-growth’ species and four ‘winter-green’ species. Among the ‘winter-green’ species, Festuca ovina and Deschampsia cespitosa were the dominant species in the sites. The ‘winter-growth’ species grew new leaves or ramets or transitioned to reproductive growth. Gentiana spathulifolia even flowered in winter. ‘Winter-growth’ made important contributions to the annual AGB, e.g. winter growth of G. spathulifolia accounted for 23.26 % of its annual AGB, while 14.74 % of the annual AGB of G. crassuloides was from winter growth. In addition, winter warming and snowfall reduction under global climate change on the eastern QTP may decrease the AGB increment of the ‘winter-growth’ and delay the green-up onset date of ‘winter-green’ species. Also, winter warming and snowfall reduction may advance the first flowering date of ‘winter-growth’ species. CONCLUSIONS: In contrast to previous views that plants on the QTP were generally considered to remain dormant in winter, our study revealed that alpine meadow plants had strong winter growth which suggested the importance of re-evaluating the dynamics of ecosystem function of alpine meadow, including its contribution to the global carbon balance. It was also shown that soil moisture availability is more important than warmer temperature in controlling the green-up onset of ‘winter-green’ species on the eastern QTP, which contrasts with the traditional view that warmer winters could advance green-up. As snowmelt is the only source of soil water in winter, the prediction of the green-up trend may be further complicated by snowfall variation in winter.

Glaciation and mountain orogeny have generated new ecologic opportunities for plants, favoring an increase in the speciation rate. Moreover, they also act as corridors or barriers for plant lineages and populations. High genetic diversity ensures that species are able to survive and adapt. Gene flow is one of the most important determinants of the genetic diversity and structure of out-crossed species, and it is easily affected by biotic and abiotic factors. The aim of this study was to characterize the genetic diversity and structure of an alpine species, Festuca ovina L., in Xingjiang, China. A total of 100 individuals from 10 populations were analyzed using six amplified fragment length polymorphism (AFLP) primer pairs. A total of 583 clear bands were generated, of which 392 were polymorphic; thus, the percentage of polymorphic bands (PPB) was 67.24%. The total and average genetic diversities were 0.2722 and 0.2006 (0.1686-0.2225), respectively. The unweighted group method with arithmetic mean (UPGMA) tree, principal coordinates analysis (PCoA) and STRUCTURE analyses revealed that these populations or individuals could be clustered into two groups. The analysis of molecular variance analysis (AMOVA) suggested that most of the genetic variance existed within a population, and the genetic differentiation (Fst) among populations was 20.71%. The Shannon differentiation coefficient (G’st) among populations was 0.2350. Limited gene flow (Nm = 0.9571) was detected across all sampling sites. The Fst and Nm presented at different levels under the genetic barriers due to fragmentation. The population genetic diversity was significant relative to environmental factors such as temperature, altitude and precipitation.

The optimal provision of green roof services is largely dependent on the health of the vegetation on the green roof, which is often affected by water availability.We assessed the impact of two substrate components, brick particle size and a polyacrylamide gel additive, and a living mulch (companion planting of Sedum spp.) on the growth, physiological and visual health of newly established plugs of Festuca ovina and Linaria vulgaris during an extreme drought (25 days) in a controlled environment greenhouse. Polyacrylamide gel and large brick substrates increased both plant species' drought tolerance whilst a Sedum living mulch had no effect. Polyacrylamide gel increased the reservoir of water that plants could access during the drought, whilst large brick promoted slower and therefore more drought resistant plant growth. Overall this study shows that polyacrylamide gel and coarser particle size substrates can significantly improve the drought tolerance of green roof plants.

Mobilization of cadmium accumulated in Festuca ovina L. roots and simultaneous immobilization of this metal by soil were studied in three chambers connected into one system containing: (1) roots in an extractant solution, (2) soil in an extractant solution, and (3) extractant solution alone. Six extractants sterilized by filtration were used: 0.1 M NaNO3 (NA), 0.1 mM desferrioxamine B (NA + DFOB) and 1 mM citric acid (NA + CA) in NA, and a water extract of soil (SE) supplemented with the same compounds. SE mobilized 53% of the Cd introduced to the system with roots. The addition of DFOB or CA to SE increased Cd extraction from roots by 17%, while the same compounds introduced to NA did not change mobilization of Cd (60% efficiency). Regardless of the extractant used, mobilization of Cd from roots was about 25% lower when extraction was done in a control system without soil. The metal released from roots was gradually immobilized by the soils loaded into all systems during a 4-day incubation. Seque...

We have measured the rates of root production and death and of root respiration in situ under two grasslands along an altitudinal gradient in the northern Pennines, UK, represented by a lowland site at 171 m in an agricultural setting, and three upland sites between 480 and 845 m. One grassland was dominated by Festuca ovina and was on a brown earth soil; the other was dominated by Juncus squarrosus and Nardus stricta and occurred on a peaty gley. The natural altitudinal gradient was extended by transplantation. Although root biomass and root production (estimated using minirhizotrons) both showed pronounced seasonal peaks, there was no simple altitudinal gradient in either variable, and neither root production nor root death rate was a simple function of altitude. Increased root accumulation in summer was a function of change in the length of the growing season, not of soil temperature. Root populations in winter were similar at all sites, showing that increased production at some sites was accompanied by increased turnover, a conclusion confirmed by cohort analyses. Respiration rate, measured in the field by extracting roots and measuring respiration at field temperature in an incubator, was unrelated to temperature. The temperature sensitivity of respiration (expressed as the slope of a plot of log respiration rate against temperature) showed no simple seasonal or altitudinal pattern. Both root growth (under Festuca) and respiration rate were, however, closely related to radiation fluxes, averaged over the previous 10 days for growth and 2 days for respiration. The temperature sensitivity of respiration was a function of soil temperature at the time of measurement. These results show that root growth and the consequent input of carbon to soil in these communities is controlled by radiation flux not temperature, and that plants growing in these upland environments may acclimate strongly to low temperatures. Most carbon cycle models assume that carbon fluxes to soil are powerfully influenced by temperature, but that assumption is based largely on short-term studies and must be reassessed.

We used outdoor microcosms in order to freely manipulate three trophic levels (ladybird/aphid/grass) at two soil fertility levels (low and high). Two hypotheses were tested: (1) that top-down control is only a mechanistic factor at high soil fertility, and (2) that herbivory increases secondary plant succession by preferentially feeding on the fast-growing early-successional grasses. Plant biomass responded dramatically to the high soil fertility treatment, as did aphid numbers in the absence of ladybirds, and ladybird activity (ladybirds feeding on aphids). At low soil fertility, plant biomass was low, aphid numbers were small, and ladybird activity was minimal. Only at high soil fertility did top-down control cause a significant response to plant biomass and species composition. The two fast-growing, early-successional grasses (Poa annua and Arrhenatherum elatius) had a greater biomass in the presence of the ladybirds compared to when the ladybirds were absent, while the slow-growing, late-successional grass (Festuca ovina) suffered. The opposite was found when ladybirds were absent but aphids present. These results suggest that herbivory may increase the rate of secondary succession, but that top-down control of herbivory by carnivores may reduce the impact of herbivory in high productivity communities.

tAnthropogenic deposition of reactive nitrogen (N) has increased during the 20th century, and is con-sidered an important driver of shifts in ecosystem functions and biodiversity loss. The objective of thepresent study was to identify those ecosystem functions that best evidence a target ecosystem’s sensitiv-ity to N deposition, taking coastal heathlands as an example. We conducted a three-year field experimentin heathlands of the island Fehmarn (Baltic Sea, North Germany), which currently are subject to a back-ground deposition of 9 kg N ha−1yr−1. We experimentally applied six levels of N fertilisation (applicationof 0, 2.5, 5, 10, 20, and 50 kg N ha−1yr−1), and quantified the growth responses of different plant speciesof different life forms (dwarf shrubs, graminoids, bryophytes, lichens) as well as shifts in the C:N ratios ofplant tissue and humus horizons. For an applicability of the experimental findings (in terms of heathlandmanagement and critical load assessment) fertilisation effects on response variables were visualised bycalculating the treatment ‘effect sizes’. The current year’s shoot increment of the dominant dwarf shrubCalluna vulgaris proved to be the most sensitive indicator to N fertilisation. Shoot increment significantlyresponded to additions of ≥ 5 kg N ha−1yr−1already in the first year, whereas flower formation of Callunavulgaris increased only in the high-N treatments. Similarly, tissue C:N ratios of vascular plants (Callunavulgaris and the graminoids Carex arenaria and Festuca ovina agg.) only decreased in the highest N treat-ments (50 and 20 kg N ha−1yr−1, respectively). In contrast, tissue C:N ratios of cryptogams respondedmore quickly and sensitively than vascular plants. For example, Cladonia spp. tissue C:N ratios respondedto N additions ≥ 5 kg N ha−1yr−1in the second study year. After three years we observed an increase incover of graminoids and a corresponding decrease of cryptogams at N fertilisation rates of ≥ 10 kg N ha−1yr−1. Soil C:N ratios proved to be an inappropriate indicator for N fertilisation at least within our three-year study period. Although current critical N loads for heathlands (10−20 kg N ha−1yr−1) were confirmedin our experiment, the immediate and highly sensitive response of the current year’s shoots of Callunavulgaris suggests that at least some ecosystem functions (e.g. dwarf shrub growth) also might respond tolow (i.e. < 10 kg N ha−1yr−1) but chronic inputs of N. Anthropogenic deposition of reactive nitrogen (N) has increased during the 20th century, and is considered an important driver of shifts in ecosystem functions and biodiversity loss. The objective of the present study was to identify those ecosystem functions that best evidence a target ecosystem’s sensitivity to N deposition, taking coastal heathlands as an example. We conducted a three-year field experiment in heathlands of the island Fehmarn (Baltic Sea, North Germany), which currently are subject to a background deposition of 9 kg N ha−1 yr−1. We experimentally applied six levels of N fertilisation (application of 0, 2.5, 5, 10, 20, and 50 kg N ha−1 yr−1), and quantified the growth responses of different plant species of different life forms (dwarf shrubs, graminoids, bryophytes, lichens) as well as shifts in the C:N ratios of plant tissue and humus horizons. For an applicability of the experimental findings (in terms of heathland management and critical load assessment) fertilisation effects on response variables were visualised by calculating the treatment ‘effect sizes’. The current year’s shoot increment of the dominant dwarf shrub Calluna vulgaris proved to be the most sensitive indicator to N fertilisation. Shoot increment significantly responded to additions of ≥ 5 kg N ha−1 yr−1 already in the first year, whereas flower formation of Calluna vulgaris increased only in the high-N treatments. Similarly, tissue C:N ratios of vascular plants (Calluna vulgaris and the graminoids Carex arenaria and Festuca ovina agg.) only decreased in the highest N treatments (50 and 20 kg N ha−1 yr−1, respectively). In contrast, tissue C:N ratios of cryptogams responded more quickly and sensitively than vascular plants. For example, Cladonia spp. tissue C:N ratios responded to N additions ≥ 5 kg N ha−1 yr−1 in the second study year. After three years we observed an increase in cover of graminoids and a corresponding decrease of cryptogams at N fertilisation rates of ≥ 10 kg N ha−1 yr−1. Soil C:N ratios proved to be an inappropriate indicator for N fertilisation at least within our three-year study period. Although current critical N loads for heathlands (10−20 kg N ha−1 yr−1) were confirmed in our experiment, the immediate and highly sensitive response of the current year’s shoots of Calluna vulgaris suggests that at least some ecosystem functions (e.g. dwarf shrub growth) also might respond to low (i.e. < 10 kg N ha−1 yr−1) but chronic inputs of N.

Complete vineyard floor cover crops were evaluated in a long-term study for their ability to regulate excessive vegetative growth of the grapevine (Vitis vinifera L.) variety ‘Cabernet Sauvignon’. Treatments were: tall fescue (Festuca arundinacea Shreb.) ‘KY-31′ and ‘Elite II’, hard fescue (Festuca ovina L.) ‘Aurora Gold’, perennial ryegrass (Lolium perenne L.), orchardgrass (Dactylis glomerata L.), and an under-trellis herbicide strip combined with KY-31 fescue interrows. Compared to herbicide-treated soil strip, Elite II fescue reduced vine pruning weights (kg/vine) 28%, individual cane weight (g) 20%, and canopy leaf layer number 25%. KY-31 fescue produced the greatest biomass and stand density, while perennial ryegrass produced the least biomass and Aurora Gold hard fescue produced the lowest stand density. Elite II fescue produced less biomass but equivalent stand density compared to KY-31 fescue. Treatments minimally impacted vine water potential (Ψmd, Ψstem), indicating that the grasses were not overly competitive with grapevines for soil moisture. All grass treatments tended to depress grapevine nitrogen levels relative to the under-trellis herbicide strip treatment, but no treatment resulted in vine nitrogen levels below the acceptable sufficiency range. Because of its establishment and growth characteristics, desired suppression of vine vegetative growth, and its low impact on crop yield, we considered Elite II fescue the optimal cover crop evaluated.

The most frequent way to establish turfgrass in the climatic conditions of the Czech Republic is sowing during spring period. The seedlings are endangered by lack of soil moisture and by high temperatures if irrigation is no available. One of the possibilities how to increase probability of successful turfgrass establishment in warm areas which suffer by low amount of precipitation is autumn term of sowing. Eight grass species and subspecies and two legumes were sown in three autumn terms (September, October and November) during 2009 and 2010. The shortest time for seedling emergence was found in average of all terms at Medicago lupulina and Trifolium repens (9 days). Lolium perenne (14) and Festuca arundinacea (17) were the fastest emerged grasses. On the other hand the longest emergence time was realised at Poa pratensis (23), Festuca rubra ssp. trichophyla (22) and Festuca ovina (21). Plants sown in September emerged in 10 days, in October 12 and in November in 29 days. September term was suitable for all used species but November term underwent most successfully Festuca rubra ssp. commutata (23% soil cover in April next year) and Lolium perenne (19%). November term was not acceptable for legumes.

Anthropogenic disturbance in alpine ecosystems is on the increase, and knowledge is needed about restoration methods and their long-term effects in these environments. Sowing seeds from introduced grass species has been a common approach to encourage the growth of new vegetation, but there is disagreement over the long-term effects. Little is known about native alternatives and their ability to promote establishment of native vegetation. We compared vegetation cover, species richness and soil conditions in a long-term field experiment in seeded sites and unseeded reference sites 21 years after seeding with a commercial seed mixture. We also tested germination and establishment of an introduced grass Festuca rubra and its native congener Festuca ovina and their effect on the establishment of the native shrub Betula nana in a greenhouse experiment in different soil types. In the long-term field experiment, the introduced Festuca species outcompeted rather than facilitated natural vegetation recovery in the seeded sites. Total vegetation cover was significantly higher in the seeded sites, but native vegetation cover and species richness were significantly higher in the unseeded sites. In the greenhouse experiment, the native F. ovina showed a tendency to facilitate the seedlings of B. nana in contrast to the introduced F. rubra , which clearly suppressed B. nana seedlings. However, seedlings of the native shrub B. nana performed best when seeded without any of the species that are presumed to be facilitators or nurse species. Suppression by seeded grass was stronger on commercial peat soil compared to fine mineral soil and coarse soil. When seeding is used in restoration efforts, it is important to choose species that do not compete strongly with the natural vegetation. Fertilization and other soil improvements may even favor strong seeded competitors at the expense of native species. Hence the combination of seeding introduced species and improving the soil may delay successful alpine natural vegetation recovery.

A regional description of the pioneer psammophilous grasslands in Poland is given. As a conseąuence of the syntaxonomic revision of 1055 phytosociological records (numerical classification according to the TWINSPAN programme) the occurrence of a single association Spergulo morisonii-Corynephoretum canescentis R. Tx. ex K. Czyżewska hoc loco is stated in Poland, and — within that association — the following syntaxa: subass. typicum subass. nova (var. typicum var. nova, var. with Artemisia campestris subsp. campestris vaf. nova, var. with Hypochoeris radicata var. nova, and intermediate form), subass. thymetosum serpyll subass. nova (var. typicum var. nova, var. with Artemisia campestris subsp. campestris var. nova, var. with Festuca ovina var. nova, and var. with F. psammophila), subass. cladinetosum mitis R. Tx. (1928) 1937 em. K. Czyżewska (var. typicum var. nova, var. with Thymus serpyllum var. nova, and var. with Polytrichum piliferum var. nova — the later and earlier phases), as well as subatlantic and subcontinental races. Nomenclatoric types are given for all the units concerned.