Aims Positive interactions are defined as non-trophic interactions where at least one of the interacting species is benefited in terms of fitness and the other remains unaffected. Nevertheless, the bidirectional feedbacks between species may be positive, neutral or negative. Thus, if facilitated species induce negative effects on their 'nurses', the assumed definition of positive interactions could be reconsidered.
Methods We assessed if ecological interactions between cushions of Azorella madreporica and their facilitated species are positive. Specifically, we tested if cover of facilitated species has any costs for cushion plants from an ecophysiological perspective, and if these costs increase with the amount of cover of facilitated species. In addition, through pathway analysis and correlations, we assessed if cover and richness of facilitated species have a direct and/or indirect effect on the fitness of cushion plants.
Important findings We found that facilitated plant species induced a significant cost for their nurses (cushion plants), and this cost increases with cover of the facilitated species. Additionally, the facilitated species exert a strong direct negative effect on the cushion's fitness and a moderate indirect negative cost evident through the nutrient status and physiological performance of cushion plants. We thus contribute evidence that positive interactions between high mountain cushion plants of central Chile and their 'facilitated' species may be an artifact more than a fact, especially when bidirectional effects are considered; contrasting with the majority of studies that document only one side of the interaction.

Aims Deserts are one of the ecosystems most sensitive to global climate change. However, there are few studies examining how changing abiotic and biotic factors under climate change will affect plant species diversity in the temperate deserts of Asia. This study aimed to: (i) characterize species distributions and diversity patterns in an Asian temperate desert; and (ii) to quantify the effects of spatial and environment variables on plant species diversity.
Methods We surveyed 61 sites to examine the relationship between plant species diversity and several spatial/environmental variables in the Gurbantunggut Desert. Spatial and environmental variables were used to predict plant species diversity in separate multiple regression and ordination models. Variation in species responses to spatial and environmental conditions was partitioned by combining these variables in a redundancy analysis (RDA) and by creating multivariate regression trees (MRT).
Important findings We found 92 plant species across the 61 sites. Elevation and geographic location were the dominant environmental factors underlying variation in site species richness. A RDA indicated that 93% of the variance in the species–environment relationships was explained by altitude, latitude, longitude, precipitation and slope position. Precipitation and topographic heterogeneity, through their effects on water availability, were more important than soil chemistry in determining the distribution of species. MRT analyses categorized communities into four groups based on latitude, soil pH and elevation, explaining 42.3% of the standardized species variance. Soil pH strongly influenced community composition within homogeneous geographic areas. Our findings suggest that precipitation and topographic heterogeneity, rather than edaphic heterogeneity, are more closely correlated to the number of species and their distributions in the temperate desert.

Aims The introduction of potentially invasive species through ornamental cultivation or for rehabilitation purposes is a serious environmental problem. They cause damage to biodiversity through loss, increased mortality or ' in situ ' selection phenomena in natural flora. Spartium junceum is a Mediterranean shrub that is not native in most areas of the Iberian Peninsula, although it is extensively grown for the rehabilitation of roadsides. We have investigated the effect on the native vegetation of an old S. junceum (Fabaceae) plantation in a conservation area in the centre of the Iberian Peninsula: the Cuenca Alta del Manzanares nature reserve in Madrid.
Methods We compared S. junceum stands with the native nanophanerophytic Cistus ladanifer community at different ecosystem properties: soil properties, temporal soil seed bank contents, standing vegetation and net primary production of annual grasslands growing in these shrublands.
Important findings The results highlighted S. junceum 's ability to become established in the new environment (marginal areas of the nature reserve) and ensure its successful growth. This is more apparent in northern and eastern exposures where this formation contacts with the core of the best conserved native vegetation in the nature reserve. Soils under Spartium showed a higher nitrogen content, indicating its capacity—in common with other legumes—to fix N, and conferring an advantage over Cistus, which is N-limited. Other soil nutrients such as phosphorus, magnesium and calcium and water availability are also higher in Spartium soils than in Cistus. Phosphorus is usually a constraint for N-fixers, but in our study, it is the most significant soil variable in both shrub formations and is important to the success of Spartium. Water availability is a key factor for Mediterranean vegetation, and particularly in autumn when soils are recharged. The Spartium formation is able to retain water as its growth produces a closer canopy than Cistus, thereby preventing water evaporation and contributing to the success of this species. Perennials are more frequent in the standing vegetation than in the seed bank, whereas therophytes are similar. The standing vegetation has therophytes and chamaephytes as the predominant growth forms in Spartium sites, and hemicryptophytes and phanerophytes in Cistus. Therophytes are dominant in Spartium and Cistus seed banks, although the first formation has more species. Spartium has a higher number of hemicryptophytes and Cistus is higher in phanerophytes. Northern and eastern aspects show significant differences in richness—with a predominance of annual weed species in Spartium —and in above-ground net primary production, probably as a consequence of the nutrients present in the soils. Ruderal annual species under Spartium (Bromus tectorum, Chenopodium album) have higher germination rates in the greenhouse than in the standing vegetation, suggesting they are at potential risk if environmental conditions change.

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.
Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.
Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.

Aims The plants of Mediterranean sea cliff ecosystems are resistant to several environmental challenges. In this study, six species typical of the coastal rocky cliffs have been analyzed in order to evaluate their diverse morphological and physiological responses to their environment across the seasons, and to examine the strategy of the ecological group to which each species belongs. Since these species are widespread across the Mediterranean region, our aim was also to highlight their ecophysiological features in habitats where the direct influence of the sea is stronger.
Methods The selected species are characteristic of the sea cliffs of Elba island (Tyrrhenian sea, Italy): the halophyte Crithmum maritimum, the semideciduous Helichrysum italicum and Lavandula stoechas and the sclerophylls Myrtus communis, Quercus ilex and Rhamnus alaternus. Four morphological traits—canopy height, leaf area, specific leaf area and leaf dry matter content—and two physiological traits—leaf water potential (LWP) and photosynthetic efficiency (PE), measured before the dawn and at midday—were analyzed. Water potential was measured by a pressure chamber and photosynthetic efficiency was determined by the analysis of chlorophyll fluorescence. Plant performance was also evaluated by calculating chronic (PI chr) and dynamic photoinhibition (PI dyn).
Important findings Crithmum maritimum showed high resistance to the recurrent dry periods, because of the high water storage capacity of its leaves and its PE declined markedly only in July, under the harshest climatic conditions. Semideciduous taxa utilize primarily an avoidance strategy, which aims at reducing the overall leaf surface, while sclerophylls mostly show a tolerance strategy towards the prevailing stressors, as demonstrated by LWP and PE, that are lower in the sclerophylls than in the semideciduous taxa during summer, due to osmoregulation and photoinhibition, respectively. Furthermore, variability of physiological parameters was higher in the sclerophylls than in the semideciduous taxa, because the former had to withstand wider oscillations of their LWP and PE. The sclerophyllous taxa underwent a slight loss of PE also in winter, likely owing to the combined action of low temperature and high irradiance. In Mediterranean sea cliff ecosystems, the stressful combination of high irradiance, high temperatures and low rainfall typical of the summer season may have been intensified by the shallow soil which displays a poor water storage capacity. On the other hand, winter stress, caused by high solar radiation and low temperatures, does not seem to seriously affect the performance of the studied species.

Aims Conversion of secondary forests to pure larch plantations is a common management practice driven by the increasing demand for timber production in Northeast China, resulting in a reduction in soil nutrient availability after a certain number of years following conversion. Nutrient resorption prior to leaf senescence was related to soil fertility, an important nutrient conservation strategy for plants, being especially significant in nutrient-poor habitats. However, the seasonal dynamics of leaf nutrients and nutrient resorption in response to secondary forest conversion to larch plantations is not well understood.
Methods A comparative experiment between larch plantations (Larix spp.) and adjacent secondary forests (dominant tree species including Quercus mongolica, Acer mono, Juglans mandshurica and Fraxinus rhynchophylla) was conducted. We examined the variations in leaf nutrient (macronutrients: N, P, K, Ca and Mg; micronutrients: Cu and Zn) concentrations of these tree species during the growing season from May to October in 2013. Nutrient resorption efficiency and proficiency were compared between Larix spp. and the broadleaved species in the secondary forests.
Important findings Results show that the seasonal variation of nutrient concentrations in leaves generally exhibited two trends, one was a downward trend for N, P, K, Cu and Zn, and another was an upward trend for Ca and Mg. The variations in foliar nutrient concentrations were mainly controlled by the developmental stage of leaves rather than by tree species. Resorption of the observed seven elements varied among the five tree species during leaf senescence. Nutrient resorption efficiency varied 6–75% of N, P, K, Mg, Cu and Zn, while Ca was not retranslocated in the senescing leaves of all species, and Mg was not retranslocated in Larix spp. Generally, Larix spp. tended to be more efficient and proficient (higher than 6–30% and 2–271% of nutrient resorption efficiency and resorption proficiency, respectively) in resorbing nutrients than the broadleaved species in the secondary forests, indicating that larch plantations had higher leaf nutrient resorption and thus nutrient use efficiency. Compared with Larix spp., more nutrients would remain in the leaf litter of the secondary forests, indicating an advantage of secondary forests in sustaining soil fertility. In contrast, the larch plantation would reuse internal nutrients rather than lose nutrients with litter fall and thus produce a positive feedback to soil nutrient availability. In summary, our results suggest that conversion from secondary forests to pure larch plantations would alter nutrient cycling through a plant-mediated pathway.

Aims Light requirements for cactus seed germination have been considered to be associated with their adult plant height and seed mass, but this has not been thoroughly studied for other succulent species. In order to understand seed photosensitivity from desert species belonging to Asparagaceae (subfamily Agavoideae) and Cactaceae, we performed a germination experiment with and without light for 12 species and 2 varieties from 1 species from the Southern Chihuahuan Desert. We also determined if adult growth is totally determined by seedling 'growth form' in cacti.
Methods We performed a germination experiment using light and darkness for 13 species from Southern Chihuahuan Desert: 10 rosette species (Asparagaceae), as well as 1 globose, 1 columnar and 2 varieties from 1 depressed-globose species (Cactaceae). The response variables were seed germination percentage and relative light germination (RLG). In addition, in order to determine if adult-globose cacti could have cylindrical seedlings, we calculated the shape index (height/width ratio) for Coryphanta clavata and Mammillaria compressa .
Important findings All species were considered neutral photoblastic. Eleven species had similar seed germination in both light and dark conditions, and three taxa (M. compressa and the two varieties of Ferocactus latispinus) showed higher germination with light than without it. Agave salmiana, M. compressa and the two varieties of F. latispinus had higher RLG than the other species. Seed mass was an important factor because with higher seed mass there was lower dependence to light. These findings support the hypothesis that small seed mass and light requirements have coevolved as an adaptation to ensure germination. One adult-globose cactus species, M. compressa, and one adult-columnar species, C. clavata, had small seeds and neutral fotoblasticism. Seedlings from these two species exposed to light were cylindrical and those under darkness conditions were columnar. Perhaps seeds from this species are able to germinate in the dark because they produce columnar seedlings with the ability to emerge from greater soil depths where sunlight cannot penetrate.

Aims Plants of similar life forms and closely related species have been observed to create similar types of plant–soil feedbacks (PSFs). However, investigations of the consistency of PSFs within species have not yielded clear results. For example, it has been reported that species create different types of PSFs in their native and introduced ranges. The aim of this project is to examine if four species create similar PSF types from soils collected from widely distributed areas within their introduced range. The soil for this project was collected from three areas in western North America. With this design, we aim to determine species- and site-specific ability to create PSFs and if the type of PSF created is consistent in all soil from all three collection areas. The species examined are Agropyron cristatum, Centaurea solstitialis, Poa pratensis and Taeniatherum caput-medusae.
Methods We used three-field collected soils (from northern Nevada, western Montana and eastern Montana) in a two-phase greenhouse experiment to quantify the type of PSFs created by four invasive species. The first phase was a conditioning phase wherein each invasive species created species-specific changes to the soil. The second phase of the experiment was the response phase wherein both the conditioning species and a native phytometer were grown in the conditioned soil and in unconditioned (control) soil. The final aboveground biomass was used to evaluate the effect of conditioning and to determine the type of PSF created by each invasive species.
Important findings Our results suggest that three of our four study species did show consistency in relation to PSF. Two species A. cristatum and T. caput-medusae consistently created PSF types that benefit conspecifics more than heterospecifics (and thus are 'invasive' PSF types) and P. pratensis consistently exhibited no, or 'neutral', feedbacks. The fourth species (C. solstitialis) was inconsistent: in one soil, no feedback was created; in other soil, an invasive PSF was created and in the last soil, a feedback that relatively benefited the native phytometer was created. Thus, PSFs appear to uniformly contribute to the success of two species (A. cristatum and T. caput-medusae) but not C. solstitialis nor P. pratensis.

Aims Biodiversity–ecosystem function experiments can test for causal relationships between planting diversity and community productivity. Planting diversity is routinely introduced as a design element in created wetlands, yet substantive support for the finding that early diversity positively affects ecosystem functioning is lacking for wetlands. We conducted a 2-year diversity–productivity experiment using freshwater wetland mesocosms to investigate community biomass production as affected by planted macrophyte functional richness.
Methods A richness gradient of macrophytes in four emergent wetland plant functional groups was established in freshwater mesocosms for two consecutive years. Species-specific aboveground morphological traits of plant size were measured at peak growth in both years; rooting depth was measured for each species in the second year. Aboveground biomass (AGB) and belowground biomass (BGB) were harvested after peak growth in the second year; first year AGB was estimated from morphological traits in constructed regression equations. Net richness effects (i.e. both complementarity effects and selection effects) were calculated using an additive partitioning method.
Important findings Species richness had a positive effect on community AGB relative to monocultures in the first year. In the second year, mean AGB was significantly reduced by competition in the most species-rich mixtures and all mixtures underyielded relative to the average monoculture. Competition for soil resources was weaker belowground, whereby root distribution at depths>20cm was reduced at the highest richness levels but overall BGB production was not affected. Changes in species biomass were strongly reflected by variation in species morphological traits, and species above and belowground performances were highly correlated. The obligate annual (Eleocharis obtusa), a dominant competitor, significantly contributed to the depression of perennial species' growth in the second growing season. To foster primary productivity with macrophyte richness in early successional communities of created wetlands where ruderal strategies are favored and competition may be stronger than species complementarity, unsystematic planting designs such as clustering the same or similar species could provide protection for some individuals. Additionally, engineering design elements fostering spatial or temporal environmental variability (e.g. microtopography) in newly created wetlands helps diversify the responses of wetland macrophyte species to their environment and could allow for greater complementarity in biomass production.

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 (GA 3) 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 GA 3 content and N:C ratios. GA 3 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 GA 3 concentrations after nutrient addition. These results indicated that differential responses of GA 3 concentrations and N:C ratios to fertilization were related to the inequality in plant heights among species.

Aims Both high and low densities of macrophyte vegetation can impair its ecosystem service function. Harvesting is often applied to macrophyte vegetation to maintain an appropriate density. Vegetation harvesting has occasionally gone awry and caused catastrophes, such as vegetation disappearance and cyanobacterial dominance in waterways and lakes. Because water depth influences macrophyte density at all life stages, the simultaneous influences of harvesting and water depth should be carefully examined. Thus, this study aims to quantify the effects of differently harvesting Elodea nuttallii on its growth and reproduction at different water depths in field experiments.
Methods Four harvest intensities (harvesting E. nuttallii plant heights equal to 25%, 50%, 75% and 100% of the water depth) were applied to E. nuttallii growing at four different water depths (60, 90, 120 and 150cm). Plant length and root length were measured. The node number, root number of each plant and number of floating plants were counted before harvesting. The harvested plant were dried to a constant weight for dry weight determination.
Important findings The rate of increase in the length and shoot number of E. nuttallii varied from ?0.012 to 0.440 day^-1 and from ?0.020 to 0.639 day^-1, respectively. Water depth>150cm would limit E. nuttallii growth. Elodea nuttallii responded to increasing water depths and low-intensity harvesting by increasing internodal length and decreasing shoot number. The larger internodal length of E. nuttallii observed in relatively deeper water was also induced by the physical strain generated by its buoyancy as its specific gravity was less than water's. The physical mechanism of removing the plant canopy by harvesting decreased E. nuttallii buoyancy and prevented floating. Harvesting increased plant production in shallow waters <90cm deep. Moreover, it is also necessary to perform three medium-intensity harvests at a water depth of 120cm and one low-intensity harvest or no harvesting at a water depth of 150cm to achieve longer lifetimes and less biomass near the water surface when the plants reach or approach the water surface.

Aims Distyly has been regarded as an adaptation to improve compatible pollination between two floral morphs with reciprocal herkogamy. The hypothesis that the different positions of anthers and stigmas within flowers as well as their reciprocal position between morphs, reduce the probability of self pollination raised by Darwin has been rarely tested. In this study, we measured stigmatic pollen loads in response to reduced reciprocal herkogamy in two Primula species.
Methods To see whether reciprocal herkogamy can increase compatible and/or reduce incompatible pollen deposition, thus promoting compatible pollination, we shortened the distance between anthers and stigmas within the flowers by changing the position of the corolla tube, to which the anthers were fused, i.e. reduced herkogamy in natural populations of Primula secundiflora and P. poissonii and quantified stigmatic pollen loads in the field over 2 years.
Important findings In both species, stigmatic pollen loads were significantly higher in the long-styled (L-morph) than in the short-styled morph (S-morph) in both control and manipulated flowers, but percentage of compatible pollen in S-morph were higher. Flowers manipulated to halve the anther–stigma distance showed a similar pattern for 2 years: total pollen grain counts on stigmas did not differ significantly but compatible pollen grains in L- and S-morphs were significantly decreased in both species. The percentage of compatible pollen loads was decreased by 68.7% in P. secundiflora and 65.3% in P. poissonii in L-morphs, while it decreased by 30.6% and 2.9% in S-morphs, respectively. Our manipulation of the relative position of anthers and stigmas in the two distylous species indicated that a lower degree of herkogamy reduced compatible but incompatible pollen transfer was likely to increase. The higher proportion of compatible pollen in the S-morph than in the L-morph in the two Primula species could be attributed to the accessibility of two-level sexual organs, floral orientations and pollinator behaviors. This is a first attempt to manipulate intraflower herkogamy for understanding adaptation of heterostyly, shedding insights into how the reciprocal herkogamy promotes compatible pollination.

Aims The aim of this article is 3-fold. First, we present an updated version of a published megaphylogeny of vascular plants that can be used in studies of plant ecology and biogeography. Second, we develop a tool that can be used by botanists and plant ecologists to generate phylogenetic hypotheses in three scenarios. Third, we use a set of regional assemblages of angiosperm trees in North America as a model system to evaluate the effect of differences in phylogenies generated using the three scenarios on the quantification of phylogenetic properties and the relationship between measures of phylogenetic properties and environment.
Methods The taxonomy and nomenclature of plant species in the megaphylogeny were standardized according to The Plant List (version 1.1). A tool for generating phylogenies was created using the R language. The robustness of derived phylogenies was evaluated using correlation and regression analyses.
Important findings An updated megaphylogeny of vascular plants (PhytoPhylo) and a tool for reconstructing phylogenies of seed plants (S.PhyloMaker) were generated. Our study shows that phylogenies generated by S.PhyloMaker using the PhytoPhylo megaphylogeny as a backbone are nearly as good as phylogeny resolved at the species level when using derived phylogenies to quantify phylogenetic properties (e.g. phylogenetic diversity and phylogenetic relatedness) of biological assemblages, and that S.PhyloMaker-generated phylogenies are robust for studies of community ecology and biogeography, particularly those seeking for patterns of phylogenetic properties along environmental gradients.