Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

How generalists coexist: the role of floral phenotype and spatial factors in the pollination systems of two Ranunculus species

Aims Competition for pollinators between phenotypically similar flowers is believed to play an important role in floral trait diversification in the angiosperms. However, in many plant communities, species with apparently similar floral phenotypes and generalist pollination systems co-flower. Here, the pollination systems of Ranunculus acris L. and Ranunculus repens L. were investigated to determine the factors enabling the species to coexist within apparently overlapping pollination niches.Methods Sympatrically flowering populations of R. acris and R. repens were investigated at three study sites in West Wales. The floral phenotypes of the two species were compared using measurements of floral morphology and spectral analyses of petal reflectance, using principal component analysis and bee and fly colour-space models. Evidence of inter-specific discrimination by foraging insects was tested for in the field and using floral arrays. The relative roles of behavioural constancy and spatial patchiness in maintaining pollinator fidelity were estimated.Important findings The floral phenotypes of R. acris and R. repens differed significantly. Social bees were highly constant when foraging at flowers of the two species and patchy floral distribution explained some of the observed fidelity. Dipterans visiting mixed floral arrays appeared to discriminate between the species, visiting more R. acris than R. repens flowers, but there was no difference in the number of visits to single-species arrays. Social bees were more likely to display constancy to flowers of R. repens in the field. Patchiness in floral distribution, subtle differences in floral phenotype, pollinator preferences and behavioural constancy are all likely to contribute to the continued coexistence of R. acris and R. repens, despite apparent overlap in their pollination niches. Such differences have the potential to facilitate the maintenance of species diversity in plant communities, even where plants appear to share similar floral phenotypes.     

Genetic diversity of Ranunculus acris L. (Ranunculaceae) populations in relation to species diversity and habitat type in grassland communities

Correlates between genetic diversity at intra- and interpopulation levels and the species diversity in plant communities are rarely investigated. Such correlates may give insights into the effect of local selective forces across different communities on the genetic diversity of local plant populations. This study has employed amplified fragment length polymorphism to assess the genetic diversity within and between 10 populations of Ranunculus acris in relation to the species diversity (richness and evenness) of grassland communities of two different habitat types, 'seminatural' and 'agriculturally improved', located in central Germany. Within-population genetic diversity estimated by Nei's unbiased gene diversity (HE) was high (0.258-0.334), and was not correlated with species richness (Pearson's r = -0.17; P = 0.64) or species evenness (Pearson's r = 0.15; P = 0.68) of the plant communities. However, the genetic differentiation between R. acris populations was significantly correlated with the difference in species evenness (Mantel's r = 0.62, P = 0.02), but not with difference in species richness of plant communities (r = -0.17, P = 0.22). Moreover, we also found that populations of R. acris from the 'seminatural' habitat were genetically different (amova, P < 0.05) from those in 'agriculturally improved' habitats, suggesting that gene flow between these habitat types is limited. The results reported in this study may indicate that habitat characteristics influence the genetic diversity of plant species.     

Vegetation dynamics following trampling in grassland and heathland in Sølendet Nature Reserve, a boreal upland area in Central Norway

Trampling is recognised as a cause of disturbance in grassland and heathland vegetation along two public nature trails in Sølendet nature reserve in central Norway. The effects of trampling by visitors and of controlled trampling were monitored over a period of 5–7 years in permanent plots. A retrogressive development was recorded as vegetation cover and height were reduced and a substantial loss of species richness and biomass took place. Some species, mainly graminoids such as Agrostis capillaris and Carex vaginata , initially appeared to be more tolerant, but were eventually reduced as well. Low, geophytic and prostrate habits of growth and hardy leaf tissues seemed to be beneficial for tolerance to trampling. Moist grassland and lichen-dominated heathland were more vulnerable to trampling than dry grassland. Even though 350–1500 visitors walked through the permanent plots along the trails each summer, effects on the soil surface remained moderate and only a minor furrow was visible on the ground.     

Effect of plant species richness on invasibility of experimental plant communities

Invasion of unsown species to artificially created assemblages of grassland species was investigated in a 3-year field experiment. In the experiment, assemblages varying in species richness (1, 2, 4, 8, and 16 species) and functional group richness (1–4, grasses, legumes, rosette forbs, and creeping forbs) were grown in control and fertilized plots, without any attempt to prevent the invasion of unsown species or to weed them. The relationship between species and functional group diversity and above-ground biomass was positive for sown species in all study years (2003, 2004 and 2005). In the latter 2 years, weed invader biomass decreased significantly with increasing biomass of sown species and their functional group richness, but not with number of species. However, no suppressive effect of species or functional group richness beyond that by increased biomass of residents was found. In fact, slight but significant positive partial effect of species richness was found, suggesting that the negative effect of the same amount of resident biomass on invaders is stronger when the biomass is composed of fewer species. The negative relationship between the number of functional groups of residents and invader biomass suggested that better coverage of functional trait space could be a mechanism promoting the resistance to invasion. In Addition, species composition of invaders were significantly related to initial composition of sown residents.     

Plant community diversity and composition affect individual plant performance

Effects of plant community diversity on ecosystem processes have recently received major attention. In contrast, effects of species richness and functional richness on individual plant performance, and their magnitude relative to effects of community composition, have been largely neglected. Therefore, we examined height, aboveground biomass, and inflorescence production of individual plants of all species present in 82 large plots of the Jena Experiment, a large grassland biodiversity experiment in Germany. These plots differed in species richness (1–60), functional richness (1–4), and community composition. On average, in more species-rich communities, plant individuals grew taller, but weighed less, were less likely to flower, and had fewer inflorescences. In plots containing legumes, non-legumes were higher and weighed more than in plots without legumes. In plots containing grasses, non-grasses were less likely to flower than in plots without grasses. This indicates that legumes positively and grasses negatively affected the performance of other species. Species richness and functional richness effects differed systematically between functional groups. The magnitude of the increase in plant height with increasing species richness was greatest in grasses and was progressively smaller in legumes, small herbs, and tall herbs. Individual aboveground biomass responses to increasing species richness also differed among functional groups and were positive for legumes, less pronouncedly positive for grasses, negative for small herbs, and more pronouncedly negative for tall herbs. Moreover, these effects of species richness differed strongly between species within these functional groups. We conclude that individual plant performance largely depends on the diversity of the surrounding community, and that the direction and magnitude of the effects of species richness and functional richness differs largely between species. Our study suggests that diversity of the surrounding community needs to be taken into account when interpreting drivers of the performance of individual plants.     

放牧干扰下高寒草甸物种、生活型和功能群多样性与生物量的关系

为明晰放牧强度对高寒草甸物种、生活型以及功能群多样性和生物量的时间累积作用以及对多样性与生物量之间相互关系的影响。选择青藏高原东北缘高寒草甸,在6个放牧强度样地连续4 a(2012—2015)进行物种、生活型、功能群多样性和生物量调查。采用重复测量方差分析和线性回归法分析放牧强度和放牧年份对物种、生活型、功能群多样性以及多样性与生物量之间的相关关系的影响。结果表明:(1)放牧强度和放牧年份均对生物量产生显著影响。随放牧强度增加,生物量显著降低。但随放牧年份延长,不同放牧强度区生物量无一致的变化规律。(2)放牧年份对物种、生活型和功能群丰富度、均匀度和优势度的影响均达到显著水平。但放牧强度和放牧年份只对物种丰富度产生交互作用,放牧强度对物种丰富度的影响具有时间的累积效应。(3)放牧干扰下不同层次多样性,仅物种丰富度与所有多样性指数显著相关,物种丰富度可以作为物种多样性测度的代表性指标。(4)多样性与生物量关系的研究,对丰富度而言,仅低放牧强度样地(Plot2)中物种丰富度随生物量增加而显著降低,其余放牧强度样地的物种、生活型和功能群丰富度均与生物量无关。对均匀度而言,高放牧强度样地(Plot4、Plot5、Plot6)生活型均匀度随生物量的增加而显著降低。对优势度而言,高放牧强度样地生活型优势度随生物量的增加而显著增加。生活型多样性可作为放牧干扰下生物量变化快速预测的有效指标。     

The response of arbuscular mycorrhizae to fertilization, mowing, and removal of dominant species in a diverse oligotrophic wet meadow

In a wet oligotrophic meadow located in the Czech Republic, a factorial experiment with treatments consisting of fertilization, mowing, and removal of the dominant species (Molinia caerulea) was established in 1994. In 1997 Holcus lanatus, Molinia caerulea, Potentilla erecta, Prunella vulgaris, and Ranunculus auricomus were examined for arbuscular mycorrhizal (AM) hyphae, arbuscles, and vesicles three times over the season. Time had a significant effect on AM in all five species. Except for Molinia arbuscles, a modal effect occurred, with the second sampling having a greater level of AM structures than the first and the third. Fertilization had the greatest effect on AM levels by decreasing the level of Holcus hyphae and vesicles, Potentilla vesicles, Prunella hyphae, and Ranunculus hyphae and vesicles. Mowing significantly increased the number of Potentilla vesicles, and the removal of dominant species had no significant effects. Interactions between time and treatments were common. Significant effects to the arbuscle:vesicle ratio were infrequent, and those that occurred were related to changes over the season. Seasonal effects appear to have a more powerful effect on AM structures and the arbuscle:vesicle ratio than do treatment effects. In a second experiment, Ranunculus auricomus, R. acris, and R. nemorosus, sampled four times over the season, showed significant changes in AM colonization. Overall, AM structures either declined over the season or increased from April to May and then declined. There was no AM colonization response to a spring fertilization in the three species. It is postulated that the patterns observed are due to phosphorus availability and seasonal changes in soil moisture and rates of root growth and turnover.     

Plant community and tissue chemistry responses to fertilizer and litter nutrient manipulations in a temperate grassland

Human-mediated nutrient amendments have widespread effects on plant communities. One of the major consequences has been the loss of species diversity under increased nutrient inputs. The loss of species can be functional group dependent with certain functional groups being more prone to decline than others. We present results from the sixth year of a long-term fertilization and litter manipulation study in an old-field grassland. We measured plant tissue chemistry (C:N ratio) to understand the role of plant physiological responses in the increase or decline of functional groups under nutrient manipulations. Fertilized plots had significantly more total aboveground biomass and live biomass than unfertilized plots, which was largely due to greater productivity by exotic C₃ grasses. We found that both fertilization and litter treatments affected plant species richness. Species richness was lower on plots that were fertilized or had litter intact; species losses were primarily from forbs and non-Poaceae graminoids. C₃ grasses and forbs had lower C:N ratios under fertilization with forbs having marginally greater %N responses to fertilization than grasses. Tissue chemistry in the C₃ grasses also varied depending on tissue type with reproductive tillers having higher C:N ratios than vegetative tillers. Although forbs had greater tissue chemistry responses to fertilization, they did not have a similar positive response in productivity and the number of forb species is decreasing on our experimental plots. Overall, differential nutrient uptake and use among functional groups influenced biomass production and species interactions, favoring exotic C₃ grasses and leading to their dominance. These data suggest functional groups may differ in their responses to anthropogenic nutrient amendments, ultimately influencing plant community composition.     

A plant diversity×water chemistry experiment in subalpine grassland

Plant diversity has been shown to drive important ecosystem functions such as productivity. At the same time, plant diversity and species composition are altered in alpine ecosystems by human impacts such as skiing. Therefore, we investigated impacts of decreased species richness and ski piste treatments on ecosystem functions in subalpine grassland.Species richness manipulations were combined with nutrient input from snow cover treated with snow additives that are commonly used on ski pistes. Three different species richness levels containing 1, 3 or 9 species randomly selected from a larger pool plus unmanipulated meadow plots were treated with four water types to simulate melt water. One water type contained the snow additive ammonium nitrate. Invasion into the communities was prevented by weeding during 2 years and allowed in three subsequent years.Higher species richness increased plant cover and biomass and decreased their variation. The number of functional groups in a plant assemblage had a positive effect on plant growth. Ammonium nitrate strongly increased biomass and plant cover after a single application but decreased species richness in originally diverse meadow plots. There was no significant interaction between species richness and water-type treatments.After the cessation of weeding, the species richness of different plot types converged within 3 years due to invasion. Nevertheless, relationships between initial species richness and plant cover remained positive.The results suggest that the diversity and species composition of alpine vegetation are important factors influencing cover and biomass, in particular during re-colonization of bare ground after disturbances such as ski-piste construction. In slow-growing alpine vegetation, initially positive diversity effects may remain even after successional convergence of species richness due to invasion. The negative effect of ammonium nitrate on species richness suggests the snow additives should only be used with care.     

Vegetation dynamics following resource manipulations in herb-rich woodland

Changes in species richness, turnover, composition and above-ground biomass of herb-rich woodland were documented following fertilizer application and water addition over three growing seasons. Addition of fertilizer significantly reduced species richness relative to unmanipulated control and water addition plots after 3 years. This change coincided with significant increases in biomass, which were largely due to increased growth of exotic annual grasses. The reductions in richness observed in the fertilized plots were a consequence of both lower rates of local colonization and enhanced rates of local extinction of the resident species. Species loss was not random; native species were lost after nutrient addition whereas exotic species were not. Nutrient limitation was more important for species coexistence in these communities than was water availability.     

Pathogenicity of Sclerotinia sclerotiorum on Ranunculus acris in Dairy Pasture

Fifty-four isolates of Sclerotinia sclerotiorum from Ranunculus acris and other natural hosts were applied as mycelial infested kibbled wheat onto 6 month-old R. acris plants in two glasshouse screening experiments. Most isolates (90%) did not differ in their pathogenicity towards R. acris. One isolate, S. sclerotiorum G45, was selected based on its ability to cause severe disease and suppress regeneration of R. acris. A field experiment was conducted to determine the efficacy of S. sclerotiorum (G45) against R. acris in infested dairy pastures in the Takaka Valley, Golden Bay, New Zealand. Isolate G45 was formulated as a wettable powder and was applied as a slurry at 20 and 40 ml/plant in December 1995. After 10 weeks, regeneration from the crown of treated plants was apparent and a second application of S. sclerotiorum was made in February 1996. Best control of R. acris was obtained when the plants were inoculated in full flower in December. At the first time of treatment, the 40 ml application of S. sclerotiorum slurry reduced the total dry weight of R. acris by an average of 57%. The second application had no effect on total dry weight, possibly because moisture levels were not sufficient for S. sclerotiorum infection. This study confirmed S. sclerotiorum to be an aggressive pathogen of R. acris under both glasshouse and field conditions. As a result, this pathogen has potential as a mycoherbicide for R. acris. Further experiments are required to explore ways of enhancing the efficacy of S. sclerotiorum against R. acris by manipulation of the host, pathogen and environment.     

Across species‐pool aggregation alters grassland productivity and diversity

Plant performance is determined by the balance of intra‐ and interspecific neighbors within an individual's zone of influence. If individuals interact over smaller scales than the scales at which communities are measured, then altering neighborhood interactions may fundamentally affect community responses. These interactions can be altered by changing the number (species richness), abundances (species evenness), and positions (species pattern) of the resident plant species, and we aimed to test whether aggregating species at planting would alter effects of species richness and evenness on biomass production at a common scale of observation in grasslands. We varied plant species richness (2, 4, or 8 species and monocultures), evenness (0.64, 0.8, or 1.0), and pattern (planted randomly or aggregated in groups of four individuals) within 1 × 1 m plots established with transplants from a pool of 16 tallgrass prairie species and assessed plot‐scale biomass production and diversity over the first three growing seasons. As expected, more species‐rich plots produced more biomass by the end of the third growing season, an effect associated with a shift from selection to complementarity effects over time. Aggregating conspecifics at a 0.25‐m scale marginally reduced biomass production across all treatments and increased diversity in the most even plots, but did not alter biodiversity effects or richness–productivity relationships. Results support the hypothesis that fine‐scale species aggregation affects diversity by promoting species coexistence in this system. However, results indicate that inherent changes in species neighborhood relationships along grassland diversity gradients may only minimally affect community (meter) – scale responses among similarly designed biodiversity–ecosystem function studies. Given that species varied in their responses to local aggregation, it may be possible to use such species‐specific results to spatially design larger‐scale grassland communities to achieve desired diversity and productivity responses.     

Local and regional processes in low-productive mountain plant communities: the roles of seed and microsite limitation in relation to grazing

It is becoming widely accepted that plant community structure is determined not only by local scale factors, but that regional factors may play considerable role. The research studying the associated processes in different environments with different species assemblages is still limited. We conducted a two-year seed sowing experiment to test whether a plant community in a low-productive mountain snowbed is limited by seed or microsite availability and how these variables depend on natural grazing. In a factorial design, half of the plots received a mixture of seeds of fourteen species naturally occurring at the study site and above ground biomass was removed from half of the plots. These treatments were applied to plots with long term grazer exclosures and to plots accessible to grazers. Both sowing and biomass removal increased the number of seedlings, the species richness of seedlings and total species richness. The number of seedlings was higher in open plots than in exclosures in the second year. Both seedling richness and total species richness were higher in open plots. Seedling recruitment was negatively related to the amount of above ground biomass and positively to the initial species richness. These results suggest that even fairly low-productive environments can be both seed and microsite limited and that these depend on grazing pressure. Natural grazing by mammal herbivores (e.g. lemmings and reindeer) favours species colonization and seedling emergence. Low-productive mountain snowbeds are prone to colonization from the local species pool and even high species richness may not constrain ingression of new species.     

Plant community responses to long-term fertilization: changes in functional group abundance drive changes in species richness

Declines in species richness due to fertilization are typically rapid and associated with increases in aboveground production. However, in a long-term experiment examining the impacts of fertilization in an early successional community, we found it took 14 years for plant species richness to significantly decline in fertilized plots, despite fertilization causing a rapid increase in aboveground production. To determine what accounted for this lag in the species richness response, we examined several potential mechanisms. We found evidence suggesting the abundance of one functional group—tall species with long-distance (runner) clonality—drove changes in species richness, and we found little support for other mechanisms. Tall runner species initially increased in abundance due to fertilization, then declined dramatically and were not abundant again until later in the experiment, when species richness and the combined biomass of all other functional groups (non-tall runner) declined. Over 86 % of the species found throughout the course of our study are non-tall runner, and there is a strong negative relationship between non-tall runner and tall runner biomass. We therefore suggest that declines in species richness in the fertilized treatment are due to high tall runner abundance that decreases the abundance and richness of non-tall runner species. By identifying the functional group that drives declines in richness due to fertilization, our results help to elucidate how fertilization decreases plant richness and also suggest that declines in richness due to fertilization can be lessened by controlling the abundance of species with a tall runner growth form.     

践踏干扰对碧塔海高寒草甸植被茎叶性状、物种多样性和功能多样性的影响

高寒草甸具有重要的生态服务功能,然而固有脆弱性使其极易遭受气候变化和人为干扰等多重因素的影响。作为滇西北旅游资源中重要的组分之一,高寒草甸吸引了大批游客前往开展徒步旅行活动,但伴随着的践踏干扰作用会不可避免地对高寒草甸生态系统带来负面影响。然而,目前关注践踏干扰对滇西北高寒草甸植被的影响,特别是植被功能性状和功能多样性如何发生变化方面的研究还十分欠缺。以云南省香格里拉市碧塔海自然保护区内的典型高寒草甸生态系统为研究对象,采用实验践踏的方式(一共5种不同强度的践踏处理)来模拟旅游活动对草甸植被的干扰作用,并以草甸植被的茎叶性状特征为切入点,重点探讨践踏干扰对茎叶性状的平均大小和变异程度的影响,以及物种丰富度(以物种形态分类为基础)和功能丰富度(以功能性状为基础)之间的关系。研究结果显示,随着践踏强度的增加,植株高度和叶片大小的平均值,而不是茎叶性状的变异程度,出现明显下降趋势。此外,物种丰富度和功能丰富度均随践踏强度的增强而减小,且两者之间呈现显著正相关关系。然而,较之轻度践踏实验组,重度践踏实验组中的功能均匀度和功能分离度水平均有所增加,表明践踏干扰可能会在短期内打破优势种对资源的绝对占有格局和减少物种间的生态位重叠程度。尽管高寒草甸对人类践踏活动有一定的承受能力,但气候变化和人为干扰等多重因素势必会改变和影响高寒草甸群落的结构和功能可持续性,这也对高寒草甸的保护与管理工作提出了更加紧迫的要求。     

Seasonality of root fungal colonization in low-alpine herbs

Arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungal colonization of Alchemilla glomerulans, Carex vaginata, Ranunculus acris ssp. pumilus and Trollius europaeus growing in low-alpine meadows in the Finnish subarctic were studied at different times during the growing season. Fungal colonization was correlated to soil soluble phosphorus (P) concentration. The influence of flower bud removal on fungal colonization was investigated in A. glomerulans, C. vaginata and R. acris and the correlation between AM and DSE colonization was studied. The fungal colonization patterns were found to be species-specific. R. acris maintained a relatively high rate of fungal colonization throughout the summer, while the rates of colonization of T. europaeus were lower and decreased towards the end of the season. A. glomerulans had constant arbuscular and vesicular colonization throughout the summer, but hyphal and DSE colonization declined towards the end of the season. C. vaginata did not form arbuscular mycorrhiza, but was colonized by DSE fungi and hyaline septate hyphae throughout the season. The soil soluble P concentration showed some seasonal variation, but was also highly variable between the study sites. Bud removal decreased arbuscular colonization of R. acris, but no unique effects were seen in any other parameters or the other species studied. The root fungal parameters correlated with soil P in some species at some sites, but no consistent trend was found. DSE colonization was positively correlated with root vesicular and hyphal colonization in some cases. The differences in fungal colonization parameters may be related to species-specific phenologies.     

Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient

Several biodiversity experiments have shown positive effects of species richness on aboveground biomass production, but highly variable responses of individual species. The well-known fact that the competitive ability of plant species depends on size differences among species, raises the question of effects of community species richness on small-stature subordinate species. We used experimental grasslands differing in species richness (1-60 species) and functional group richness (one to four functional groups) to study biodiversity effects on biomass production and ecophysiological traits of five small-stature herbs (Bellis perennis, Plantago media, Glechoma hederacea, Ranunculus repens and Veronica chamaedrys). We found that ecophysiological adaptations, known as typical shade-tolerance strategies, played an important role with increasing species richness and in relation to a decrease in transmitted light. Specific leaf area and leaf area ratio increased, while area-based leaf nitrogen decreased with increasing community species richness. Community species richness did not affect daily leaf carbohydrate turnover of V. chamaedrys and P. media indicating that these species maintained efficiency of photosynthesis even in low-light environments. This suggests an important possible mechanism of complementarity in such grasslands, whereby smaller species contribute to a better overall efficiency of light use. Nevertheless, these species rarely contributed a large proportion to community biomass production or achieved higher yields in mixtures than expected from monocultures. It seems likely that the allocation to aboveground plant organs to optimise carbon assimilation limited the investment in belowground organs to acquire nutrients and thus hindered these species from increasing their performance in multi-species mixtures.     

Time course of plant diversity effects on Centaurea jacea establishment and the role of competition and herbivory

Aims Invasion resistance in experimental plant communities is known to increase with increasing diversity and further to depend on the presence of particular functional groups. To test whether these effects also hold true for the invader establishment phase beyond the seedling stage, we studied survival and performance of Centaurea jacea L. (brown knapweed) planted into experimental grassland communities of varying plant biodiversity over three consecutive years. Moreover, we analysed the role of insect herbivory and biomass of the recipient community for mediating diversity effects.Methods In 2005, seedlings of Centaurea were transplanted into experimental grassland communities (the Jena Experiment) covering a species richness (1–60) and functional group richness (1–4) gradient. Half of these transplants and the community surrounding them in each plot were sprayed with insecticide while the other half served as control. In 2006 and 2007 (during the second and third year after transplantation), we recorded survival, growth-related (e.g. transplant biomass, height) and reproduction-related traits (e.g. number of flower heads). Annual data on community aboveground biomass served as covariate to investigate mediating effects of aboveground competition with the recipient community.Important findings Species richness was the most important factor responsible for Centaurea limitation. Higher levels of diversity decreased survival and all performance traits in both years. These diversity effects were partly driven by community biomass, but not fully explained by that covariate, suggesting the importance also of further processes. The influence of functional group richness was strong in the second year after transplantation and weaker in the third year. Among the particular functional groups, only the presence of legumes showed strong negative effects on Centaurea survival and weak negative effects on growth and reproduction, the latter two being mediated by biomass. Insect herbivore reduction considerably benefited Centaurea in sprayed monocultures, where it grew significantly larger than in all other diversity levels and than in the control subplots. We conclude that effects of plant community properties on invading individuals change in the course of establishment, that plant species richness effects are also important during later stages of establishment, and that biomass (especially at high diversity) and herbivory (especially at low diversity) of the recipient community are important in mediating community effects on invaders.     

Reflectance Measurements of Glossy Petals of Ranunculus lingua (Ranunculaceae) and of Non-Glossy Petals of Heliopsis helianthoides (Asteraceae)

Abstract: A spectrophotometric comparison of the reflectance of glossy petals of Ranunculus lingua (Ranunculaceae) and non-glossy petals of Heliopsis helianthoides (Asteraceae) was made in relation to the angle of incidence of the incoming light. Additionally, the reflectance of petals of R. flammula, R. acris and R. repens (Ranunculaceae) was measured for comparative purposes. In the examined Ranunculus species, gloss is present around mirror geometry over the whole spectral range measured, including the UV region where it accentuates the UV peak, and even at the basal UV-absorbing area of the petals. According to a colour hexagon evaluation of the reflectance data, these gloss phenomena diminish the colour contrast of the petal against the background. Nevertheless, due to their brightness, they may serve as an attractant or predictor for pollinators and other visiting insects, possibly acting as a beacon while approaching the flower. The position of the reflectance maximum in the UV around 340 nm is in accordance with the sensitivity maximum of the UV photoreceptor of potential pollinators like Apidae. The mean reflectance values of the four examined Ranunculus species do not show any clear correlation with their anatomical make-up