Aims Grasslands are the world's most extensive terrestrial ecosystem, and are a major feed source for livestock. Meeting increasing demand for meat and other dairy products in a sustainable manner is a big challenge. At a field scale, Global Positioning System and ground-based sensor technologies provide promising tools for grassland and herd management with high precision. With the growth in availability of spaceborne remote sensing data, it is therefore important to revisit the relevant methods and applications that can exploit this imagery. In this article, we have reviewed the (i) current status of grassland monitoring/observation methods and applications based on satellite remote sensing data, (ii) the technological and methodological developments to retrieve different grassland biophysical parameters and management characteristics (i.e. degradation, grazing intensity) and (iii) identified the key remaining challenges and some new upcoming trends for future development.
Important findings The retrieval of grassland biophysical parameters have evolved in recent years from classical regression analysis to more complex, efficient and robust modeling approaches, driven by satellite data, and are likely to continue to be the most robust method for deriving grassland information, however these require more high quality calibration and validation data. We found that the hypertemporal satellite data are widely used for time series generation, and particularly to overcome cloud contamination issues, but the current low spatial resolution of these instruments precludes their use for field-scale application in many countries. This trend may change with the current rise in launch of satellite constellations, such as RapidEye, Sentinel-2 and even the microsatellites such as those operated by Skybox Imaging. Microwave imagery has not been widely used for grassland applications, and a better understanding of the backscatter behaviour from different phenological stages is needed for more reliable products in cloudy regions. The development of hyperspectral satellite instrumentation and analytical methods will help for more detailed discrimination of habitat types, and the development of tools for greater end-user operation.

Aims Invasive species often have higher relative growth rates (RGR) than their native counterparts. Nutrient use efficiency, total leaf area and specific leaf area (SLA) are traits that may confer RGR differences between natives and invasives, but trait differences are less prominent when the invasive species belongs to the same plant functional type as the dominant native species. Here, we test if traits displayed soon after germination confer an early size advantage. Specifically, we predicted that invasive species seedlings grow faster than the natives because they lack trade-offs that more strongly constrain the growth of native species.
Methods We quantified plant morphological and physiological traits and RGR during early seedling growth at high and low nutrient levels in three dominant perennial native C₄ grasses: Panicum virgatum L. (switchgrass), Schizachyrium scoparium (Michx.) Nash (little bluestem) and Andropogon gerardii Vitman (big bluestem); and a perennial C₄ exotic invasive grass, Sorghum halepense (L.) Pers. (Johnsongrass).
Important findings After 2 weeks of growth, Johnsongrass seedlings had greater biomass, SLA and photosynthetic nitrogen use efficiency, but lower leaf N concentrations (% leaf N) and root:shoot ratio than natives. As growth continued, Johnsongrass more quickly produced larger and thicker leaves than the natives, which dampened the growth advantage past the first 2 to 3 weeks of growth. Investment in carbon gain appears to be the best explanation for the early growth advantage of Johnsongrass. In natives, growth was constrained by an apparent trade-off between allocation to root biomass, which reduced SLA, and production of leaves with high N content, which increased carbon gain. In Johnsongrass, root:shoot ratio did not interact with other traits, and % leaf N was decoupled from RGR as a result of a trade-off between the positive indirect association of % leaf N with RGR and the negative direct association of % leaf N with RGR.

Aims Alien species are commonly considered as harmful weeds capable of decreasing native biodiversity and threatening ecosystems. Despite this assumption, little is known about the long-term patterns of the native–alien relationships associated with human disturbed managed landscapes. This study aims to elucidate the community dynamics associated with a successional gradient in Chilean Mediterranean grasslands, considering both native and alien species.
Methods Species richness (natives and aliens separately) and life-form (annuals and perennials) were recorded in four Chilean post-agricultural grazed grasslands each covering a broad successional gradient (from 1 to 40 years since crop abandonment). A detrended correspondence analysis (DCA), mixed model effects analyses and correlation tests were conducted to assess how this temporal gradient influenced natives and aliens through community dynamics.
Important findings Our results show different life-form patterns between natives and aliens over time. Aliens were mainly represented by annuals (especially ruderals and weeds), which were established at the beginning of succession. Annual aliens also predominated at mid-successional stages, but in old grasslands native species were slightly more representative than alien ones within the community. In the late successional states, positive or no correlations at all between alien and native species richness suggested the absence of competition between both species groups, as a result of different strategies in occupation of the space. Community dynamics over time constitute a net gain in biodiversity, increasing natives and maintaining a general alien pool, allowing the coexistence of both. Biotic interactions including facilitation and/or tolerance processes might be occurring in Chilean post-agricultural grasslands, a fact that contradicts the accepted idea of the alien species as contenders.

Aims Species interactions regulate the invasiveness of non-native species and as declines of native tree species escalate, exotic tree species that offer supplementary resources to animal seed-dispersers should expand their distributions as they fill ecological roles. Our primary objective was to forecast impacts from an imminent biological invasion (laurel wilt disease) by quantifying resources provided by native (threatened) and exotic fruits (disease-resistant) and associated bird foraging preferences.
Methods In the southeastern USA, we tested for redundancy among the resources provided by native and exotic fruits to overwintering birds. Comparisons between abundant subcanopy species Persea borbonia (native) and Cinnamomum camphora (exotic) were paramount considering the widespread disease-induced decline of P. borbonia, and the biological and phylogenetic similarities between these species. Across two winter survey periods, we quantified fruit removal and documented bird species using motion-activated cameras in the field. Physical and chemical fruit characteristics were also quantified.
Important findings Foraging bouts on both P. borbonia and C. camphora fruits were documented for four native bird species. There was no difference in selectivity between fruit types during Year 1 of our survey, but there was a significant preference for C. camphora fruit in Year 2; the change in preference was correlated with significantly lower temperatures in Year 2. While the pulp/seed ratio and moisture content differed, the nutritional content of fruit pulp (g/100g) was similar between fruit types. Given the apparent redundancy among these native and exotic fruit resources, we forecast increases in the consumption and dispersal of exotic propagules following the widespread laurel wilt disease-induced decline of P. borbonia and other native fruit bearing members of Lauraceae. This empirically based prediction is among the first to document exotic forest pathogens as indirect threats to native bird–plant interactions and potential facilitators of exotic plant invasion.

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.
Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.
Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.

Aims Grasslands are widely used for production of livestock, which depend on the nutritive value of herbage species. However, there are still large uncertainties as to how grazing, precipitation and growing season interactively affect herbage nutritive value.
Methods Here, based on a grazing experiment with seven grazing intensities in an Inner Mongolian grassland since the year 2005, we analysed nutritive value of four dominant grass species from June to September in both a relatively wetter year (2008) and a drier year (2010). Herbage with high nutritive value is characterized by high concentration of crude protein, high concentration of cellulase digestible organic matter and low concentration of neutral detergent fibre.
Important findings We found that (i) grazing significantly increased the nutritive value of Leymus chinensis, Agropyron michnoi and Cleistogenes squarrosa but had minor effects on the nutritive value of Stipa grandis. (ii) For all species, nutritive values were greater in the wetter year than in the drier year and were greatest in the early growing season (June) and lowest at the end of the growing season (September). Inter-annual and inter-seasonal variations in nutritive value were much greater for L. chinensis and A. michnoi than for C. squarrosa and S. grandis, suggesting higher water use efficiency for the latter two species. (iii) Grazing significantly decreased the drought resistance of three species, but not of S. grandis. (iv) Grazing significantly increased inter-month variation in nutritive value for L. chinensis and A. michnoi but had relatively minor or no effects on that of C. squarrosa and S. grandis. Therefore, grazing effects on species nutritive values showed strong inter-annual and seasonal patterns, and species-specific responses might be related to species traits (i.e. water use efficiency, digestibility). Our results have important implications for ecosystem management of arid and semi-arid grasslands under intense grazing and global climatic change.

Aims and Methods We propose a standard protocol at the landscape to continental scale for examining to what extent the range of ecological conditions found in temperate latitudes explains the variations in climber species richness and traits. The protocol was tested in forests of the two Americas. The data set included 151 climber species. We selected four categorical traits and grouped these species into six clusters with regard to these traits. Floristic records of American forests were first gathered into alliances, second combined with bioclimatic indices (rainfall, temperature, continentality). We obtained a total of 59 vegetational units in which we calculated values of climber species richness and proportion of clusters. Vegetational units were ultimately gathered into five forest formations (characterized by leaf longevity). Wetlands and uplands were considered separately.
Important findings Our results emphasize clear trends in large-scale patterns of climber distribution, independently of taxonomy. Climber species richness (in particular woody climbers) peaks in moist and warm upland forests with oceanic climates, and where conifers are rare. In flooded areas, climber richness is also very high and peaks in seasonally flooded large floodplains. In ecological conditions of frost, dryness or lack of nutrients, climber species richness, abundance and trait diversity decline, resulting in the dominance of small, twining and deciduous life traits.

Aims The aim of this article is 4-fold: (i) to update species richness of bryophytes for each of the Chinese provinces based on the most current knowledge on distributions of bryophytes in China, (ii) to provide a set of analyses based on the updated species richness data and the environmental variables used in a recent article on species richness of bryophytes in China, (iii) to expand the analysis presented in the recent article by relating species richness of bryophytes to over 15 additional climatic variables and (iv) to determine the degree to which the relationships between bryophyte species richness and environmental variables that were reported in the recent article might have been biased.
Methods Over 180 literatures with national, provincial and local species lists of bryophytes in China were used in this study. Taxonomy and nomenclature of bryophytes in China were standardized according to The Plant List. Correlation and regression analyses were used to examine the relationships between species richness or species density of bryophytes in Chinese provinces and environmental variables.
Important findings On average, each Chinese province possesses 700.6 species of bryophytes, which is 112.1 species more than previously reported. With the updated species richness data reported in this study, stronger relationships between species richness of bryophytes and environmental variables have been found, compared with those found in a recently published study for China. When single environmental variables were considered, precipitation-related variables were, on average, more strongly correlated with species richness and species density than were temperature-related variables. Environmental variables were on average correlated more strongly with species density than with species richness of bryophytes at the regional scale in China. Our study showed that measures quantifying the average and variation of environmental conditions within each Chinese province explained 82.7% and 71.1% of the variation in species richness of liverworts and mosses, respectively, and explained 86.5% and 70.7% of the variation in species density of liverworts and mosses, respectively.

Aims Predicting suitable habitat distribution is an effective way to protect rare or endangered medicinal plants. Cornus officinalis is a perennial tree growing in forest edge and its air-dried pericarp is one of the traditional Chinese medicines (TCM) with significant medicinal values. In recent years, C. officinalis has undergone severe degeneration of its natural habitat owing to growing market demands and unprecedented damage to the forests. Moreover, the degeneration of suitable habitat has threatened the supply of medicinal materials, and even led to the extinction of some engendered medicinal plant species. In this case, there is a great risk to introduce and cultivate medicinal plants if planners determine the suitable cultivation regions based on personal subjective experience alone. Therefore, predicting suitable potential habitat distribution of medicinal plants (e.g. C. officinalis) and revealing the environmental factors determining such distribution patterns are important to habitat conservation and environmental restoration.
Methods In this article, we report the results of a study on the habitat distribution of C. officinalis using maximum entropy (Maxent) modeling and fuzzy logics together with loganin content and environmental variables. The localities of 106 C. officinalis in China were collected by our group and other researchers and used as occurrence data. The loganin content of 234 C. officinalis germplasm resources were tested by high-performance liquid chromatography (HPLC) and used as content data. 79 environmental variables were selected and processed with multicollinearity test by using Pearson Correlation Coefficient (r) to determine a set of independent variables. The chosen variables were then processed in the fuzzy linear model according to the cell values (maximum, minimum) of localities with estimated loganin content. The SDMtoolbox was used to spatially rarefy occurrence data and prepare bias files. Furthermore, combined Maxent modeling and fuzzy logics were used to predict the suitable habitat of C. officinalis. The modeling result was validated using null-model method.
Important findings As a result, six environmental factors including tmin3, prec3, bio4, alt, bio12 and bio3 were determined as key influential factors that mostly affected both the habitat suitability and active ingredient of C. officinalis. The highly suitable regions of C. officinalis mainly distribute in a 'core distribution zone' of the east-central China. The statistically significant AUC value indicated that combined Maxent modeling and fuzzy logics could be used to predict the suitable habitat distribution of medicinal plants. Furthermore, our results confirm that ecological factors played critical roles in assessing suitable geographical regions as well as active ingredient of plants, highlighting the need for effective habitat rehabilitation and resource conservation.

Aims In Mediterranean-type ecosystem, the Cape Fynbos, legumes may be able to switch between soil N and atmospheric N 2 sources during growth to adjust the carbon costs of N acquisition. This study investigated the utilization of different inorganic N sources by Virgilia divaricata, a native legume from the Mediterranean-type ecosystem of the Cape Floristic Region.
Methods Plants were cultivated in sterile quartz sand, supplied with 25% strength Long Ashton nutrient solution, modified to contain 500 μM Phosphate. At the phosphate level (500 μM), plants were treated with 500 μM NH 4 NO 3 (treatment named N), or grown in N-free nutrient solution and inoculated with effective Burkholderia sp. (Bact.) or treated with combined N sources (500 μM NH 4 NO 3) and inoculated with effective Burkholderia sp. (N+Bact.).
Important findings The application of NH 4 NO 3 to the legumes resulted in a greater increase in plant dry matter. Carbon construction costs were higher in plants that were supplied with mineral and symbiotic N sources. Maximum photosynthetic rates per leaf area was maintained, irrespective of the N sources. Although the plant roots were nodulated, the plant dependence on N 2 fixation decreased with addition of N. Roots and nodules of the plants solely reliant on N 2 fixation showed an increase in glutamine content. These results show that V. divaricata is highly adapted for growth at the forest margin. Fynbos and possibly anthropic soils by utilizing both atmospheric and soil N sources.

Aims The effects of clouds are now recognized as critically important to the understanding of climate change impacts on ecosystems. Regardless, few studies have focused specifically on the ecophysiological responses of plants to clouds. Most continental mountain ranges are characterized by common convective cloud formation in the afternoons, yet little is known regarding this influence on plant water and carbon relations. Here we compare the ecophysiology of two contrasting, yet ubiquitous growth forms, needle-leaf and broadleaf, under representative cloud regimes of the Snowy Range, Medicine Bow Mountains, southeastern Wyoming, USA.
Methods Photosynthetic gas exchange, water use efficiency, xylem water potentials and micrometeorological data were measured on representative clear, overcast and partly cloudy days during the summers of 2012 and 2013 for two indigenous broadleaf (Caltha leptosepala and Arnica parryi) and two needle-leaf species (Picea engelmannii and Abies lasiocarpa) that co-occur contiguously.
Important findings Reductions in sunlight with cloud cover resulted in more dramatic declines in photosynthesis for the two broadleaf species (ca. 50–70% reduction) versus the two conifers (no significant difference). In addition, the presence of clouds corresponded with lower leaf conductance, transpiration and plant water status in all species. However, the more constant photosynthesis in conifers under all cloud conditions, coupled with reduced transpiration, resulted in greater water use efficiency (ca. 25% higher) than the broadleaf species. These differences appear to implicate the potential importance of natural cloud patterns in the adaptive ecophysiology of these two contrasting, but common, plant growth forms.

Aims Relatively few studies so far have assessed how ongoing global warming will affect the photosynthetic performance of dryland plant species. We evaluated the effects of warming on the photosynthetic rates of 10 species with contrasting functional attributes, and whether their functional traits modulated photosynthetic responses to warming.
Methods A common garden experiment was conducted over 2 years with distinct environmental conditions (drier vs. wetter year). The experiment was designed as a randomized block design with two treatments: warming (control vs. ~2.9°C temperature increase) and species (Agropyron cristatum, Festuca ovina, Lygeum spartum, Medicago sativa, Plantago lanceolata, Psoralea bituminosa, Sanguisorba minor, Hedysarum coronarium, Dorycnium pentaphyllum and Phlomis herba-venti). We linked functional traits measurements with temporal variations in photosynthetic responses to warming.
Important findings In the drier year, warming increased photosynthetic rates at the beginning of the growing season, suggesting a modification in the growing period (earlier spring). In the wetter year, functional traits modulated photosynthetic responses to warming. Larger species with shorter leaves (e.g. M. sativa) had higher photosynthetic rates under warming compared to smaller species with larger leaves (e.g. F. ovina). Our results highlight the importance of (i) studying photosynthetic responses along different years and (ii) considering functional traits when evaluating photosynthetic responses to climate change, particularly in stressful environments such as drylands.

Aims Nitrogen (N) and phosphorus (P) constitute essential elements for plant growth and their availability influence species diversity in herbaceous plant communities. Legumes exhibit relatively high abundance in N-limited soils. Moreover, the legumes' N:P ratios are much higher than those of the other plant species grown in the same site, probably because they are able to fix atmospheric N 2. The objective of this study was to determine how the relative proportion in N and P availability and the restriction of legumes to fix atmospheric N 2 affect: (i) the primary productivity of plant species, (ii) species composition and (iii) N and P concentrations of species.
Methods In an outdoor experiment, mixtures containing grasses, legumes and non-legume forbs were established in 32 containers under four soil treatments (control, N addition, P addition and disinfected soil), in a completely randomized design with eight replicates. Plant growth was examined when N and P were limited in the control soil:sand mixture, in a pot experiment sown with Plantago lanceolata .
Important findings The pot experiment indicated that both N and P were limiting for the growth of P. lanceolata. Soil treatments affected primary productivity and species composition. Legumes had a relatively high abundance in the control and their growth was favoured, especially that of Medicago sativa, by P addition. Grasses' growth was increased by the addition of N. Inhibition of rhizobia resulted in poor growth of legumes and concomitant higher growth of grasses, in comparison to the control. The N:P ratios of non-legume species differed between treatments and were always higher in the legume species, even in the disinfected soil. The latter provides evidence that the high N concentrations found in legumes are a physiological characteristic of this specific group of plants.

Aims Diversity–productivity relationships among herbaceous species have mostly been studied in grasslands, while less is known about diversity effects among weedy species with a short life cycle.
Methods We studied diversity–productivity relationships, shoot density, size and allometry in experimental communities of different species richness (one, three, six, and nine species), functional group number (one to three functional groups: grasses, small herbs and tall herbs) and functional group evenness (even and uneven number of species per functional group) based on a pool of nine arable weed species with a short life cycle in a 2-year experiment.
Important findings Higher species richness increased above- and belowground biomass production in both years of the experiment. Additive partitioning showed that positive selection effects increased with increasing species richness and functional group number, while positive complementarity effects were greater when tall herbs were present. Relative yield totals were larger than 1 across all species richness levels but did not increase with species richness, which is consistent with constant positive complementarity effects. Community biomass production and diversity effects increased in the second year of the experiment, when communities achieved greater shoot densities and average shoot sizes. At the community level, varying productivity was mainly attributable to variation in mean shoot sizes. Tall herbs reached greater observed/expected relative yields (=overyielding) due to increased shoot sizes, underyielding of small herbs was mainly attributable to decreased shoot sizes, while grasses partly compensated for reduced shoot sizes by increasing densities. Shifts in community-level density–size relationships and changes in shoot allometry in favour of greater height growth indicated that a greater biomass at a given density was due to increased dimensions of occupied canopy space. We conclude that diversity effects are also possible among short-lived arable weed species, but selection effects through sizes differences among species are key for positive species richness–productivity relationships.

Aims Reintroduction has become an important tool for the management of endangered plant species. We tested the little-explored effects of small-scale environmental variation, genotypic composition (i.e. identity of genotypes) and genotypic diversity on the population survival of the regionally rare clonal plant Ranunculus reptans. For this species of periodically inundated lakeshores, genetic differentiation had been reported between populations and between short-flooded and long-flooded microsites within populations.
Methods We established 306 experimental test populations at a previously unoccupied lakeshore, comprising either monocultures of 32 genotypes, mixtures of genotypes within populations or mixtures of genotypes between populations. In 2000, three years after planting out at the experimental site, a long-lasting flood caused the death of half of the experimental populations. In 2003, an extreme drought resulted in the lowest summer water levels ever measured.
Important findings Despite these climatic extremes, 27% of the established populations survived in December 2003. The success of experimental populations largely differed between microsites. Moreover, the success of genotype monocultures depended on genotype and source population. Genetic differentiation between microsites played a minor role for the success of reintroduction. After the flood, populations planted with genotypes from different source populations increased in abundance, whereas populations with genotypes from single source populations and genotype monocultures decreased. In 2014, several small patches of Ranunculus clones were still present, but plants were strongly intermingled, which precluded their assignment to the original treatments. We conclude that sources for reintroductions need to be selected carefully. Moreover, mixtures of plants from different populations appear to be the best choice for successful reintroduction, at least in unpredictably varying environments.