Aims The majority of angiosperms are pollinated by animals, and this interaction is of enormous importance in both agricultural and natural systems. Pollinator behavior is influenced by plants' floral traits, and these traits may be modified by interactions with other community members. In recent years, knowledge of ecological linkages between above- and belowground organisms has grown tremendously. Soil communities are extremely diverse, and when their interactions with plants influence floral characteristics, they have the potential to alter pollinator attraction and visitation, but plant–pollinator interactions have been neglected in studies of the direct and indirect effects of soil organism–root interactions. Here, we review these belowground interactions, focusing on the effects of nitrogen-fixing bacteria, arbuscular mycorrhizal fungi and root-feeding herbivores, and their effects on floral traits and pollinators. Further, we identify gaps in our knowledge of these indirect effects and recommend promising directions and topics that should be addressed by future research.
Important findings Belowground organisms can influence a wide variety of floral traits that are important mediators of pollinator attraction, including the number and size of flowers and nectar and pollen production. Other traits that are known to influence pollinators in some plant species, such as floral volatiles, color and nectar composition, have rarely or never been examined in the context of belowground plant interactions. Despite clear effects on flowers, relatively few studies have measured pollinator responses to belowground interactions. When these indirect effects have been studied, both arbuscular mycorrhizal fungi and root herbivores were found to shift pollinator visitation patterns. Depending on the interaction, these changes may either increase or decrease pollinator attraction. Finally, we discuss future directions for ecological studies that will more fully integrate belowground ecology with pollination biology. We advocate a multilevel approach to these questions to not only document indirect effect pathways between soil interactions and pollination but also identify the mechanisms driving changes in pollinator impacts and the resultant effects on plant fitness. A more thorough understanding of these indirect interactions will advance ecological theory and may inform management strategies in agriculture and conservation biology.

Aims Measures of plot-to-plot phylogenetic dissimilarity and beta diversity are providing a powerful tool for understanding the complex ecological and evolutionary mechanisms that drive community assembly.
Methods Here, we review the properties of some previously published dissimilarity measures that are based on minimum or average phylogenetic dissimilarity between species in different plots.
Important findings We first show that some of these measures violate the basic condition that for two identical plots the measures take the value zero. They also violate the condition that the dissimilarity between two identical plots should always be lower than that between two different plots. Such erratic behavior renders these measures unsuitable for measuring plot-to-plot phylogenetic dissimilarity. We next propose a new measure that satisfies these conditions, thus providing a more reasonable way for measuring phylogenetic dissimilarity.

Aims Understanding the relationships among disturbance, invasion and species change is essential for effective management of many systems. We investigated relationships among fire history, invasion by a native tree species, Allocasuarina huegeliana, and diversity change to understand the potential drivers of plant community alteration in a complex and biodiverse system.
Methods We used plant species surveys from 1983 and 2011 to quantify species loss/gain and thence compositional changes. Additionally, we surveyed population densities of the invasive species and collated long-term fire history data for each site. General linear models and non-parametric models were used to assess the strength of relationships between the three variables of interest.
Important findings Within the last 30 years, ~11% of the plant species richness was lost from the reserve. At an individual site level, we found only a 4% average decrease in overall plant species richness, but large species losses and gains that imply considerable compositional shifts. Though such shifts might be expected over 30 years, many of the gained species were common, potentially opportunistic species, while those lost were often locally rare woody perennials. In addition, gained species tended to be expanding their recorded range westward suggesting that they may be responding to the regional drying climate. The relationship between invasion density and species loss was strong over all spatial scales. We identified a potential state change to dominance by the native invasive particularly as high densities prevented species gain at the site scale. In these extreme cases of high invasive density and high biodiversity loss, we argue that there may be a need to directly address the expanding native population.

Aims Grassland is the most widely distributed vegetation type on the Xizang Plateau. Accurate remote sensing estimation of the grassland aboveground biomass (AGB) in this region is influenced by the types of vegetation indexes (VIs) used, the grain size (resolution) of the remote sensing data and the targeted ecosystem features. This study attempts to answer the following questions: (i) Which VI can most accurately reflect the grassland AGB distribution on the Xizang Plateau? (ii) How does the grain size of remote sensing imagery affect AGB reflection? (iii) What is the spatial distribution pattern of the grassland AGB on the plateau and its relationship with the climate?
Methods We investigated 90 sample sites and measured site-specific AGBs using the harvest method for three grassland types (alpine meadow, alpine steppe and desert steppe). For each sample site, four VIs, namely, Normalized Difference VI (NDVI), Enhanced VI, Normalized Difference Water Index (NDWI) and Modified Soil-Adjusted VI (MSAVI) were extracted from the Moderate Resolution Imaging Spectroradiometer (MODIS) products with grain sizes of 250 m and 1 km. Linear regression models were employed to identify the best estimator of the AGB for the entire grassland and the three individual grassland types. Paired Wilcoxon tests were applied to assess the grain size effect on the AGB estimation. General linear models were used to quantify the relationships between the spatial distribution of the grassland AGB and climatic factors.
Important findings The results showed that the best estimator for the entire grassland AGB on the Xizang Plateau was MSAVI at a 250 m grain size (MSAVI 250 m). For each individual grassland type, the best estimator was MSAVI at a grain size of 250 m for alpine meadow, NDWI at a grain size of 1 km for alpine steppe and NDVI at a grain size of 1 km for desert steppe. The explanation ability of each VI for the grassland AGB did not significantly differ for the two grain sizes. Based on the best fit model (AGB =^-10.80 + 139.13 MSAVI 250 m), the spatial pattern of the grassland AGB on the plateau was characterized. The AGB varied from 1 to 136g m ?2. Approximately 59% of total spatial variation in the AGB for the entire grassland was explained by the combination of the mean annual precipitation (MAP) and mean annual temperature. The explanatory power of MAP was weaker for each individual grassland type than that for the entire grassland. This study illustrated the high efficiency of the VIs derived from MODIS data in the grassland AGB estimation on the Xizang Plateau due to the vegetation homogeneity within a 1×1 km pixel in this region. Furthermore, MAP is a primary driver on the spatial variation of AGB at a regional scale.

Aims The Inclusive Niche Hypothesis has not been validated for plants using ecophysiological performance. The few experiments have measured growth and competition but not the physiological response of plants. A metallophyte plant that hyperaccumulates aluminium (Al), Plantago almogravensis, showed a defined spatial distribution by occurring mostly on vegetation gaps associated with metalliferous areas (geochemical islands). This case was used to determine, in situ, whether the Inclusive Niche Hypothesis was suitable to explain the extent of the species realized niche.
Methods The vegetation associated with P. almogravensis geochemical islands in the SW coast of Portugal was mapped. The biotic (neighbouring plants) and abiotic (edaphic) components of the niche were correlated with parameters of the plant's ecological and physiological performances (plant density and cover; leaf C and N concentration and isotopic composition; growth). The results were obtained using image analysis, abundance and morphological measures, isotopic signatures and chemical composition.
Important findings The species showed better physiological performance where its ecological performance was lower due to trade-offs with environmental constraints. The species' realized niche was mostly limited by shrub competition and soil Al-toxicity. These limits contribute to explain the rarity status of the species: the species has a poor capacity to compete but, due to an enhanced Al-tolerance and Al-hyperaccumulator trait, has the ability to find refuge in geochemical islands that are too harsh for most other species. This work, based on ecophysiological field studies, provides support for the Inclusive Niche Hypothesis relating to plant species.

Aims Soil CO₂ emission from steppes is affected by soil properties and vegetation in different successional stages. Primary and secondary succession of plants frequently occurred at the meadow steppe in Songnen Plain, Northeast China, which indicates the large uncertainty associated with CO₂ emission in this environment. This study aims to investigate the temporal variations of soil respiration (Rs) and the effect of plant succession on cumulative soil CO₂ emission during the growing season.
Methods Using a LI-6400 soil CO₂ flux system, Rs of five vegetation types which represented different stages of plant succession in meadow steppes of Songnen Plain, China, was investigated during the growing seasons of 2011 and 2012.
Important findings Soil temperature (Ts) was the dominant controlling factor of Rs, which could explain ~64% of the change in CO₂ fluxes. The Q 10 values of Rs were ranged from 2.0 to 6.7, showing a decreasing trend with the plant successional stages. The cumulative CO₂ emission increased with the degree of vegetation succession and it averaged to 316±6g C m ?2 (ranges: 74.8±6.7 to 516.5±11.4g C m ?2) during the growing season. The magnitude of soil CO₂ emission during the growing season was positively correlated with aboveground plant biomass, soil organic carbon content and mean soil water content, while negatively linked to mean Ts, pH, electrical conductivity and exchangeable sodium percentages. The results implied that soil CO₂ emission increased with the development of plant communities toward more advanced stages. Our findings provided valuable information for understanding the variations of CO₂ emission in the process of vegetation succession.

Aims To clarify whether variation in leaf traits with climate differs with scale, i.e. across species and within a species, and to detect whether plant functional group affects species-specific response.
Methods Leaf dry matter content (LDMC), specific leaf area (SLA), mass- and area-based leaf N (N mass, N area) and leaf P concentrations (P mass, P area) and leaf chlorophyll concentration (SPAD) were measured for 92 woody plant species in two botanical gardens in China. The two gardens share plant species in common but differ in climate. Leaf trait variation between the two gardens was examined via mean comparison at three scales: all species together, species grouped into plant functional groups and within a species. A meta-analysis was performed to summarize the species-specific responses.
Important findings At the scale of all species together, LDMC, SLA, P mass and N mass were significantly lower in the dry-cold habitat than in the wet-warm one, whereas N area and SPAD showed an inverse pattern, indicating a significant environmental effect. The meta-analysis showed that the above-mentioned patterns persisted for SLA, N area and SPAD but not for the other variables at the species-specific scale, indicating that intraspecific variation affects the overall pattern of LDMC, P mass and N mass and P area. In terms of species-specific response, positive, negative or nonsignificant patterns were observed among the 92 species. Contrary to our prediction, species-specific responses within a functional group were not statistically more similar than those among functional groups. Our results indicated that leaf trait variation captured climatic difference yet species-specific responses were quite diverse irrespective of plant functional group, providing new insights for interpreting trait variability with climate.

Aims Plant functional traits determine how plants respond to environmental factors and influence ecosystem processes. Among them, root traits and analyses of relations between above and below-ground traits in natural communities are scarce. Methods we characterized a set of above- and below-ground traits of three dominant shrub species in a semiarid shrub-steppe that had contrasting leaf phenological habits (deciduous, semideciduous and evergreen). We analysed if there was coordination among above- and below-ground resource economics patterns: i.e. patterns of biomass allocation, construction costs and lifespan.
Important findings Above- and below-ground traits and their resource economics relations pointed to species-specific functional strategies to cope with drought and poor soils and to a species ranking of fast to slow whole-plant strategies in terms of resource uptake, biomass construction costs and turnover. The deciduous shrub, Proustia cuneifolia, had relatively deep and even distribution of roots, and high proportion of short-lived tissues of low C construction costs: it had high fine to coarse root and high leaf-to-stem biomass ratios, high specific leaf area (SLA), and stems of low wood density. This strategy allows Proustia to maximize and coordinate above- and below-ground resources uptake as long as the most limiting factor (water) is available, but at the cost of having relative high plant biomass turnover. The evergreen Porlieria chilensis, instead, displayed a more conservative and slow strategy in terms of resource economics. It had ~80% of the roots in the 40cm topsoil profile, low proportion of fine compared with coarse roots and low leaf-to-stem ratios, low SLA and stems of high wood density, i.e. it invested in C costly tissues that, overall, persist longer but probably at the cost of having lower plant resource uptake rates. Traits in the semideciduous Adesmia bedwellii were in between these two functional extremes. Our results revealed high functional diversity and above- and below-ground complementarity in resource economics among these three codominant species in the Chilean coastal desert.

Aims Arid environments are resource limited, with scarcity of water the key limiting factor, but hyper-arid environments are rarely studied. We test for spatial and temporal variation in ecologically important characteristics to deduce plant adaptations to the extreme climate.
Methods The endangered Sinai Thyme (Thymus decussatus) exists as a set of patches on mountaintops within the St Katherine Protectorate, South Sinai, a hyper-arid environment with rare events of good rains (every 10–15 years).
Important findings From spatial and temporal patterns of plant mortality, size, condition and flowering among 10 patches on the Mt Sinai massif, we deduce that the incidence and amount of flowering responds relatively quickly (1–2 years) to rainfall fluctuations, but plant growth respond only very slowly. Small individuals are most at risk of death during drought, and a high proportion of plants were dead at the end of 8 years of very low or no rainfall. No recruitment of seedlings was observed even in years of good rainfall. Droughts are expected to become increasingly frequent due to climate change; this may have important consequences for Sinai Thyme and also its associated herbivores, such as the Critically Endangered Sinai Baton Blue (Pseudophilotes sinaicus) whose larval stage feeds exclusively upon the flowers of this plant.

Aims Soil heterogeneity is common in natural habitats. It may trigger foraging responses (placing more ramets and/or roots in nutrient-rich patches than in nutrient-poor patches) and further affect the growth of plants. However, the impact of soil heterogeneity on competitive interactions has been little tested.
Methods We conducted a greenhouse experiment to investigate the effects of soil heterogeneity on intraspecific competition with a stoloniferous herb Hydrocotyle vulgaris. We grew one (without competition) or nine ramets (with competition) of H. vulgaris under a homogeneous environment and two heterogeneous environments differing in patch size (large or small patches). In the heterogeneous treatment, the soil consisted of the same number of nutrient-rich and nutrient-poor patches arranged in a chessboard manner, and in the homogeneous treatment, the soil was an even mixture of the same amount of the nutrient-rich and the nutrient-poor soil.
Important findings Irrespective of intraspecific competition, H. vulgaris showed foraging responses to soil heterogeneity in the large patch treatment, e.g. it produced significantly more biomass, ramets, aboveground mass and root mass in the nutrient-rich patches than in the nutrient-poor patches. In the small patch treatment, foraging responses were observed when intraspecific competition was present, but responses were not observed when there was no competition. However, we find a significant effect of soil heterogeneity on neither overall growth nor competitive intensity of H. vulgaris. Our results suggest that foraging responses to soil heterogeneity may not necessarily be adaptive and intraspecific competition may not be influenced by soil heterogeneity.