Why are graminoid species more dominant? Trait-mediated plant-soil feedbacks shape community composition.

Species traits may determine plant interactions along with soil microbiome, further shaping plant-soil feedbacks (PSFs). However, how plant traits modulate PSFs and, consequently, the dominance of plant functional groups remains unclear. We used a combination of field surveys and a two-phase PSF experiment to investigate whether forbs and graminoids differed in PSFs and in their trait-PSF associations. When grown in forb-conditioned soils, forbs experienced stronger negative feedbacks, while graminoids experienced positive feedbacks. Graminoid-conditioned soil resulted in neutral PSFs for both functional types. Forbs with thin roots and small seeds showed more-negative PSFs than those with thick roots and large seeds. Conversely, graminoids with acquisitive root and leaf traits (i.e., thin roots and thin leaves) demonstrated greater positive PSFs than graminoids with thick roots and tough leaves. By distinguishing overall and soil biota-mediated PSFs, we found that the associations between plant traits and PSFs within both functional groups were mainly mediated by soil biota. A simulation model demonstrated that such differences in PSFs could lead to a dominance of graminoids over forbs in natural plant communities, which might explain why graminoids dominate in grasslands. Our study provides new insights into the differentiation and adaptation of plant life-history strategies under selection pressures imposed by soil biota.

Changes in microbial composition explain the contrasting responses of glucose and lignin decomposition to soil acidification in an alpine grassland.

Soil acidification often suppresses microbial growth and activities, resulting in a negative impact on soil organic carbon (C) decomposition. While the detrimental effects of acidification on soil and plant properties have been extensively studied, less attention has been paid on the shifts in soil microbial communities and their influences of the decomposition of organic C with different chemical complexities. Taking advantage of an acid addition experiment in a Tibetan alpine meadow, here we examined the response of soil microbial communities to soil acidification and microbial effect on the decomposition of organic C with different chemical complexities (i.e., glucose and lignin, representing labile and recalcitrant C respectively). We found that soil acidification had no impact on microbial respiration and microbial abundance even though it decreased bacterial diversity significantly. Soil acidification increased the relative abundance of some microbial taxa, like Alphaproteobacteria and Acidobacteriia in bacteria increased by 36 %, 284 %, and Eurotiomycetes, Sordariomycetes and Leotiomycetes in fungi increased by 145 %, 279 % and 12.7-fold, but decreased the relative abundance of Acidimicrobiia by 33 % in highest acid addition treatment. Changes in microbial communities (bacterial and fungal community composition, the diversity of bacterial community and the ratio of fungi to bacteria) are significantly related to the decomposition of glucose and lignin. More specifically, soil acidification decreased the decomposition of glucose but increased the decomposition of lignin, indicating a trade-off between the decomposition of labile and recalcitrant soil organic C under soil acidification. Overall, shifts in microbial communities under soil acidification might be accompanied by an increased ability to break down more recalcitrant C. This trade-off between the decomposition of labile and recalcitrant C may change soil C quality under future acid deposition scenarios.

Nutrient-induced acidification modulates soil biodiversity-function relationships.

Nutrient enrichment is a major global change component that often disrupts the relationship between aboveground biodiversity and ecosystem functions by promoting species dominance, altering trophic interactions, and reducing ecosystem stability. Emerging evidence indicates that nutrient enrichment also reduces soil biodiversity and weakens the relationship between belowground biodiversity and ecosystem functions, but the underlying mechanisms remain largely unclear. Here, we explore the effects of nutrient enrichment on soil properties, soil biodiversity, and multiple ecosystem functions through a 13-year field experiment. We show that soil acidification induced by nutrient enrichment, rather than changes in mineral nutrient and carbon (C) availability, is the primary factor negatively affecting the relationship between soil diversity and ecosystem multifunctionality. Nitrogen and phosphorus additions significantly reduce soil pH, diversity of bacteria, fungi and nematodes, as well as an array of ecosystem functions related to C and nutrient cycling. Effects of nutrient enrichment on microbial diversity also have negative consequences at higher trophic levels on the diversity of microbivorous nematodes. These results indicate that nutrient-induced acidification can cascade up its impacts along the soil food webs and influence ecosystem functioning, providing novel insight into the mechanisms through which nutrient enrichment influences soil community and ecosystem properties.

Losses of low-germinating, slow-growing species prevent grassland composition recovery from nutrient amendment.

Nutrient enrichment often alters the biomass and species composition of plant communities, but the extent to which these changes are reversible after the cessation of nutrient addition is not well-understood. Our 22-year experiment (15 years for nutrient addition and 7 years for recovery), conducted in an alpine meadow, showed that soil nitrogen concentration and pH recovered rapidly after cessation of nutrient addition. However, this was not accompanied by a full recovery of plant community composition. An incomplete recovery in plant diversity and a directional shift in species composition from grass dominance to forb dominance were observed 7 years after the nutrient addition ended. Strikingy, the historically dominant sedges with low germination rate and slow growth rate and nitrogen-fixing legumes with low germination rate were unable to re-establish after nutrient addition ceased. By contrast, rapid recovery of aboveground biomass was observed after nutrient cessation as the increase in forb biomass only partially compensated for the decline in grass biomass. These results indicate that anthropogenic nutrient input can have long-lasting effects on the structure, but not the soil chemistry and plant biomass, of grassland communities, and that the recovery of soil chemical properties and plant biomass does not necessarily guarantee the restoration of plant community structure. These findings have important implications for the management and recovery of grassland communities, many of which are experiencing alterations in resource input.

Nematode biomass changes along an elevational gradient are trophic group dependent but independent of body size.

Aboveground, large and higher trophic-level organisms often respond more strongly to environmental changes than small and lower trophic-level organisms. However, whether this trophic or size-dependent sensitivity also applies to the most abundant animals, microscopic soil-borne nematodes, remains largely unknown. Here, we sampled an altitudinal transect across the Tibetan Plateau and applied a community-weighted mean (CWM) approach to test how differences in climatic and edaphic properties affect nematode CWM biomass at the level of community, trophic group and taxon mean biomass within trophic groups. We found that climatic and edaphic properties, particularly soil water-related properties, positively affected nematode CWM biomass, with no overall impact of altitude on nematode CWM biomass. Higher trophic-level omnivorous and predatory nematodes responded more strongly to climatic and edaphic properties, particularly to temperature, soil pH, and soil water content than lower trophic-level bacterivorous and fungivorous nematodes. However, these differences were likely not (only) driven by size, as we did not observe significant interactions between climatic and edaphic properties and mean biomasses within trophic groups. Together, our research implies a stronger, size-independent trophic sensitivity of higher trophic-level nematodes compared with lower trophic-level ones. Therefore, our findings provide new insights into the mechanisms underlying nematode body size structure in alpine grasslands and highlight that traits independent of size need to be found to explain increased sensitivity of higher trophic-level nematodes to climatic and edaphic properties, which might affect soil functioning.

Taxonomic and functional biogeographies of soil bacterial communities across the Tibet plateau are better explained by abiotic conditions than distance and plant community composition.

The processes governing soil bacteria biogeography are still not fully understood. It remains unknown how the importance of environmental filtering and dispersal differs between bacterial taxonomic and functional biogeography, and whether their importance is scale-dependent. We sampled soils across the Tibet plateau, with distances among plots ranging from 20 m to 1550 km. Taxonomic composition of bacterial community was characterized by 16S amplicon sequencing and functional community composition by qPCR targeting 9 functional groups involved in N dynamics. Factors representing climate, soil and plant community were measured to assess different facets of environmental dissimilarity. Both bacterial taxonomic and functional dissimilarities were more related to abiotic dissimilarity than biotic (vegetation) dissimilarity or distance. Taxonomic dissimilarity was mostly explained by differences in soil pH and mean annual temperature (MAT), while functional dissimilarity was linked to differences in soil N and P availabilities and N:P ratio. Soil pH and MAT remained the main determinants of taxonomic dissimilarity across spatial scales. In contrast, the explanatory variables of N-related functional dissimilarity varied across the scales, with soil moisture and organic matter having the highest role across short distances (<~330 km), and available P, N:P ratio and distance being important over long distances (>~660 km). Our results demonstrate how biodiversity dimension (taxonomic versus functional aspects) and spatial scale influence the factors driving soil bacterial biogeography.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Can nitrogen supersede host identity in shaping the community composition of foliar endophytic fungi in an alpine meadow ecosystem?

The availability of limiting nutrients plays a crucial role in shaping communities of endophytes. Moreover, whether fungal endophytes are host-specific remains controversial. We hypothesized that in a harsh and nitrogen (N)-deficient area, diversity and community composition of foliar endophytic fungi (FEFs) varied substantially among plots with experimentally elevated levels of macronutrients, and thus, N availability, instead of host species identity, would have a greater influence in structuring fungal communities at different scales. We also expected an important subset of taxa shared among numerous host species and N gradients to form a community-wide core microbiome. We measured the leaf functional traits and community structures of FEFs of three commonly seen species in an alpine meadow nested with a long-term N fertilization experiment. We found that host plant identity was a powerful factor driving the endophytic fungal community in leaves, even in habitats where productivity was strongly limited by nitrogen (p < 0.001). We also found that within the same host, nitrogen was an important driving force for the composition of the endophytic fungi community (p < 0.05). In addition, the leaf carbon content was the most important functional trait that limited the diversity of endophytic fungi (p < 0.001). Finally, we documented a distinct core microbiome shared among our three focal species and N gradients.

N-Induced Species Loss Dampened by Clipping Mainly Through Suppressing Dominant Species in an Alpine Meadow.

Nitrogen addition and clipping can exert substantial impact on species diversity but their interactions and the underlying mechanisms still remain unclear. Resource competition theory holds that sufficiently strong competitive ability of dominant species can lead to the losses of subordinate species through competitive exclusion, while niche differentiation theory suggests that the persistence of subordinate species in competitive systems can be promoted by guaranteeing positive growth rates of rare species. Taking advantage of a field experiment with nitrogen addition (10 g N m-2 year-1) and different clipping intensities (2, 15, and 30 cm) treatments in a Tibetan alpine meadow across 2015-2020, we assessed the relative importance of competitively dominant species and niche differentiation in driving species diversity changes via using community weighted mean (CWM) and variation coefficient of nearest neighbor distance (CV_NND) of functional traits including height, specific leaf area (SLA) and leaf dry matter content (LDMC). We show that nitrogen enrichment drove a strong plant diversity loss (P < 0.001). Clipping at different intensities had little effect on species diversity, but it can reduce the N-induced diversity loss. Nitrogen addition and clipping caused changes in community diversity were mainly indirectly attributed to their effects on community functional composition, and the competitive ability of dominant species. Nitrogen increased the CWM of functional traits to improve the competitive ability of dominant species. In contrast, clipping influenced species diversity positively by decreasing CWMheight (P < 0.001), and also negatively by increasing CWMSLA (P < 0.001) and decreasing CV_NNDSLA (P < 0.05). Interacting with N addition, clipping resulted in a neutral effect on species diversity, because clipping could offset the negative effects of nitrogen addition through an opposite effect on CWMheight. This study provides new insights into the mechanisms of diversity maintenance with respect to nitrogen addition and clipping. Thus, clipping is recommended as a useful management strategy to alleviate the species loss caused by nutrients enrichment and maintain the diversity of grassland ecosystems.

Current patterns of plant diversity and phylogenetic structure on the Kunlun Mountains.

Large-scale patterns of biodiversity and the underlying mechanisms that regulate these patterns are central topics in biogeography and macroecology. The Qinghai-Tibet Plateau serves as a natural laboratory for studying these issues. However, most previous studies have focused on the entire Qinghai-Tibet Plateau, leaving independent physical geographic subunits in the region less well understood. We studied the current plant diversity of the Kunlun Mountains, an independent physical geographic subunit located in northwestern China on the northern edge of the Qinghai-Tibet Plateau. We integrated measures of species distribution, geological history, and phylogeography, and analyzed the taxonomic richness, phylogenetic diversity, and community phylogenetic structure of the current plant diversity in the area. The distribution patterns of 1911 seed plants showed that species were distributed mainly in the eastern regions of the Kunlun Mountains. The taxonomic richness, phylogenetic diversity, and genera richness showed that the eastern regions of the Kunlun Mountains should be the priority area of biodiversity conservation, particularly the southeastern regions. The proportion of Chinese endemic species inhabiting the Kunlun Mountains and their floristic similarity may indicate that the current patterns of species diversity were favored via species colonization. The Hengduan Mountains, a biodiversity hotspot, is likely the largest source of species colonization of the Kunlun Mountains after the Quaternary. The net relatedness index indicated that 20 of the 28 communities examined were phylogenetically dispersed, while the remaining communities were phylogenetically clustered. The nearest taxon index indicated that 27 of the 28 communities were phylogenetically clustered. These results suggest that species colonization and habitat filtering may have contributed to the current plant diversity of the Kunlun Mountains via ecological and evolutionary processes, and habitat filtering may play an important role in this ecological process.

Multimode Gesture Recognition Algorithm Based on Convolutional Long Short-Term Memory Network.

Gesture recognition utilizes deep learning network model to automatically extract deep features of data; however, traditional machine learning algorithms rely on manual feature extraction and poor model generalization ability. In this paper, a multimodal gesture recognition algorithm based on convolutional long-term memory network is proposed. First, a convolutional neural network (CNN) is employed to automatically extract the deeply hidden features of multimodal gesture data. Then, a time series model is constructed using a long short-term memory (LSTM) network to learn the long-term dependence of multimodal gesture features on the time series. On this basis, the classification of multimodal gestures is realized by the SoftMax classifier. Finally, the method is experimented and evaluated on two dynamic gesture datasets, VIVA and NVGesture. Experimental results indicate that the accuracy rates of the proposed method on the VIVA and NVGesture datasets are 92.55% and 87.38%, respectively, and its recognition accuracy and convergence performance are better than those of other comparison algorithms.

Light plasticity of germination on the eastern Tibetan Plateau: Phylogeny, trait, and environmental correlates.

Seeds often exhibit great plasticity of germination in response to environmental variability and uncertainty. The causes of this plasticity, however, remain poorly understood, and comparative phylogenic analyses of such plasticity are rare. Here, we analyzed a field germination dataset including 474 species exposed to three different levels of light availability, using comparative phylogenetic methods. We calculated the plasticity of germination in response to light availability (PGGP) based on the maximum germination proportion (GPmax), PGT50 based on the time required to reach 50% of GPmax, PGRGV based on the relative germination velocity (RGV), and PGTotal based on all three of these germination traits. We found that closely related species shared similar light plasticity of germination behavior. Different aspects of germination plasticity in response to light availability were related to specific traits or local environment. PGGP was associated with adult longevity and local water habitat, while PGT50 was related to seed mass and local water habitat, and PGRGV was marginally significantly related to plant height. PGTotal was significantly associated with adult longevity and water habitat. These results suggested that different aspects of germination plasticity were located at specific niche dimension, and local habitats with sufficient soil moisture induced great plasticity germination in response to light environment. As such, they can simplify our understanding of germination, promote the exploration of the general law of germination, and further increase our understanding of species diversity maintenance, adaptation, and evolution from the perspective of germination.

Differential Mechanisms Drive Species Loss Under Artificial Shade and Fertilization in the Alpine Meadow of the Tibetan Plateau.

Fertilization is an effective management strategy to promote community biomass but can simultaneously reduce species diversity in many grassland systems. Shifts in competition for resources have been proposed to explain the decline in plant species diversity due to fertilization, yet the underlying mechanism driving species loss remains controversial. This uncertainty may be driven by variation in aboveground and belowground resource availability. However, experiments simultaneously manipulating both light availability and soil nutrients are rare. Using a 6-year field experiment to manipulate light availability (via shade cloth) and soil nutrients (via fertilizer addition), we tested this resource competition hypothesis in a species-rich alpine meadow by examining the variation of species traits associated with the capacity of light acquisition within these treatments. Our results showed that artificial shade decreased community biomass accumulation whereas fertilization increased it. In contrast, both shade and fertilization reduced species diversity. Extinction of non-Gramineae species (e.g., Fabaceae and Cyperaceae) was the main reason for species diversity decline. Species loss can be explained by the limitation of light availability and predicted by species traits associated with light acquisition capability under fertilization and low light tolerance under artificial shade. Specifically, fertilization eliminated species with lower stature and artificial shade exterminated species with the higher light compensation point (LCP). The findings suggest that light availability is consistently important for plant growth and that low competitiveness for light under fertilization and intolerance of low light conditions under artificial shade trigger species loss process in the alpine meadow. Our experiment helps clarify the mechanisms of how artificial shade and fertilization decreased species diversity and highlight that LCP, which tends to be neglected by most of the studies, is one of the vital drivers in determining species coexistence.

Current biogeographical roles of the Kunlun Mountains.

Large-scale patterns of biodiversity and formation have garnered increasing attention in biogeography and macroecology. The Qinghai-Tibet Plateau (QTP) is an ideal area for exploring these issues. However, the QTP consists of multiple geographic subunits, which are understudied. The Kunlun Mountains is a geographical subunit situated in the northern edge of the QTP, in northwest China. The diversity pattern, community phylogenetic structures, and biogeographical roles of the current flora of the Kunlun Mountains were analyzed by collecting and integrating plant distribution, regional geological evolution, and phylogeography. A total of 1911 species, 397 genera, and 75 families present on the Kunlun Mountains, of which 29.8% of the seed plants were endemic to China. The mean divergence time (MDT) of the Kunlun Mountains flora was in the early Miocene (19.40 Ma). Analysis of plant diversity and MDT indicated that the eastern regions of the Kunlun Mountains were the center of species richness, endemic taxa, and ancient taxa. Geographical origins analysis showed that the Kunlun Mountains flora was diverse and that numerous clades were from East Asia and Tethyan. Analysis of geographical origins and geological history together highlighted that the extant biodiversity on the Kunlun Mountains appeared through species recolonization after climatic fluctuations and glaciations during the Quaternary. The nearest taxon index speculated that habitat filtering was the most important driving force for biodiversity patterns. These results suggest that the biogeographical roles of the Kunlun Mountains are corridor and sink, and the corresponding key processes are species extinction and immigration. The Kunlun Mountains also form a barrier, representing a boundary among multiple floras, and convert the Qinghai-Tibet Plateau into a relatively closed geographical unit.

The Complex Biodiversity-Ecosystem Function Relationships for the Qinghai-Tibetan Grassland Community.

Despite the long history of the study of the biodiversity-ecosystem function relationship, uncertainty remains about the relationship of natural grassland ecosystems under stressful conditions. Recently, trait- and phylogenetic-based tests provide a powerful way to detect the relationship in different spaces but have seldom been applied to stressful zones on a large spatial scale. We selected Qinghai-Tibetan as the study area and collected a grassland community database involving 581 communities. We calculated biomass and species', functional, and phylogenetic diversity of each community and examined their relationships by using linear and non-linear regression models. Results showed an overall positive biodiversity-productivity relationship in species', functional and phylogenetic space. The relationship, however, was non-linear, in which biodiversity explained better the variation in community biomass when species diversity was more than a threshold, showing a weak effect of biodiversity on ecosystem function in low species diversity communities. We also found a filled triangle for the limit of the relationship between species and functional diversity, implying that functional diversity differs significantly among communities when their species diversity is low but finally converges to be a constant with increasing communities' species diversity. Our study suggests that multiple niche processes may structure the grassland communities, and their forces tend to balance in high-biodiversity communities.

Direct and indirect facilitation affect community productivity through changes in functional diversity in an alpine system.

BACKGROUND AND AIMS: Facilitation is an important ecological process for plant community structure and functional composition. Although direct facilitation has accrued most of the evidence so far, indirect facilitation is ubiquitous in nature and it has an enormous potential to explain community structuring. In this study, we assess the effect of direct and indirect facilitation on community productivity via taxonomic and functional diversity. METHODS: In an alpine community on the Tibetan Plateau, we manipulated the presence of the shrub Dasiphora fruticosa and graminoids in a fenced meadow and a grazed meadow to quantify the effects of direct and indirect facilitation. We measured four plant traits: height, lateral spread, specific leaf area (SLA) and leaf dry matter content (LDMC) of forbs; calculated two metrics of functional diversity [range of trait and community-weighted mean (CWM) of trait]; and assessed the responses of functional diversity to shrub facilitation. We used structural equation modelling to explore how shrubs directly and indirectly drove community productivity via taxonomic diversity and functional diversity. KEY RESULTS: We found stronger effects from herbivore-mediated indirect facilitation than direct facilitation on productivity and taxonomic diversity, regardless of the presence of graminoids. For functional diversity, the range and CWM of height and SLA, rather than lateral spread and LDMC, generally increased due to direct and indirect facilitation. Moreover, we found that the range of traits played a primary role over taxonomic diversity and CWM of traits in terms of shrub effects on community productivity. CONCLUSIONS: Our study reveals that the mechanism of shrub direct and indirect facilitation of community productivity in this alpine community is expanding the realized niche (i.e. expanding range of traits). Our findings indicate that facilitators might increase trait dispersion in the local community, which could alleviate the effect of environmental filters on trait values in harsh environments, thereby contributing to ecosystem functioning.

High soil pH enhances the network interactions among bacterial and archaeal microbiota in alpine grasslands of the Tibetan Plateau.

Soil functions and processes are driven by complex microbial interactions. It is, therefore, critical to understand the coexistence patterns of soil microbiota, especially in fragile alpine ecosystems. We identified biogeographic patterns in the network-level topological features of the soil microbial co-occurrence network in the Tibetan alpine grasslands, based on high-throughput sequencing. We verified that soil pH was the most important environmental variable for predicting network-level topological features of soil microbial co-occurrence networks. Associations among soil microbiota were enhanced with increasing pH (5.17-8.92), and the network was the most stable at neutral pH. Moreover, node-level topological features suggested that the archaeal operational taxonomic units, compared with bacterial operational taxonomic units, hold a central role in the co-occurrence network. Network-level topological features revealed closer connections among soil microbiota in the steppe ecosystem than in the meadow ecosystem. Therefore, our study demonstrated that soil pH served as a critical environmental filter that influenced the potential associations and ecological signature of soil microbiota in the Tibetan alpine grasslands. These findings provide a new perspective on the distinct biogeographic patterns of co-occurrence networks, to explore the ecological role of soil microbiota and thus help manage soil bacterial and archaeal communities for provisioning alpine ecosystem services.

Fast and furious: Early differences in growth rate drive short-term plant dominance and exclusion under eutrophication.

The reduction of plant diversity following eutrophication threatens many ecosystems worldwide. Yet, the mechanisms by which species are lost following nutrient enrichment are still not completely understood, nor are the details of when such mechanisms act during the growing season, which hampers understanding and the development of mitigation strategies.Using a common garden competition experiment, we found that early-season differences in growth rates among five perennial grass species measured in monoculture predicted short-term competitive dominance in pairwise combinations and that the proportion of variance explained was particularly greater under a fertilization treatment.We also examined the role of early-season growth rate in determining the outcome of competition along an experimental nutrient gradient in an alpine meadow. Early differences in growth rate between species predicted short-term competitive dominance under both ambient and fertilized conditions and competitive exclusion under fertilized conditions.The results of these two studies suggest that plant species growing faster during the early stage of the growing season gain a competitive advantage over species that initially grow more slowly, and that this advantage is magnified under fertilization. This finding is consistent with the theory of asymmetric competition for light in which fast-growing species can intercept incident light and hence outcompete and exclude slower-growing (and hence shorter) species. We predict that the current chronic nutrient inputs into many terrestrial ecosystems worldwide will reduce plant diversity and maintain a low biodiversity state by continuously favoring fast-growing species. Biodiversity management strategies should focus on controlling nutrient inputs and reducing the growth of fast-growing species early in the season.

Phenological variation of flower longevity and duration of sex phases in a protandrous alpine plant: potential causes and fitness significance.

BACKGROUND: Flower longevity plays an important role in pollen dispersal and reproductive success in plants. In dichogamous plants, the duration of anthesis as well as the time allocated to male and female functions can vary in response to intrinsic factors (e.g., flowering time and resource allocation) and pollination context along a growth season. However, the fitness consequences of phenological dynamics have rarely been examined. This study aims to unravel the potential causes driving variation in flower longevity, duration of sex phases, and phenotypic sex during a flowering season of strongly protandrous Aconitum gymnandrum, and particularly reproductive consequences of the phenological pattern. RESULTS: Population floral sex ratio shifted from completely male at the beginning to completely female at the end of the season, as is common in other protandrous plants. Phenological dynamics of the floral sex ratio and the duration of sex phases caused a shift from femaleness to maleness in the mean phenotypic sex over the whole season. Floral longevity was negatively correlated with flower size and positively affected by temperature. Early flowers within inflorescences rather than early-flowering individuals emphasized the duration of female over male phase. Owing to the dominance of male-phase flowers, early flowering for individual flowers and plants, or female-biased sex resulted in higher pollen deposition per flower and seed set. At the flower level, flower longevity positively affected female reproductive success, while the effect of flower size was negative. By contrast, plant-level female reproductive success was negatively affected by flower longevity but positively correlated to flower size. CONCLUSIONS: The major result of this study lies in elucidating the relationship between variation in phenological sex expression and floral longevity and their fitness consequences of protandrous A. gymnandrum. The contrasting results on female fitness for individual flowers and plants contribute to our current understanding of the adaptive significance of floral longevity.

Direct and indirect effects of temperature and precipitation on alpine seed banks in the Tibetan Plateau.

Plant community responses to global environmental change focus primarily on aboveground vegetation; however, the important role of the seed bank is frequently neglected. Specifically, the direct and indirect effects of changes in temperature and precipitation on seed banks remain poorly understood, yet seed banks provide a vital source of ecosystem resilience to global environmental change. We used a structural equation model to explore the direct and indirect effects of temperature, precipitation, and other biotic and abiotic factors on soil seed bank community composition using 1,026 soil seed bank samples from 57 sites along an elevation gradient that served as a space-for-time substitution for changing climate in the Tibetan Plateau. Seed bank richness was negatively correlated with both precipitation and temperature, but neither climate factor affected seed bank density. Temperature was also negatively correlated with vegetation species richness, which was positively correlated with seed bank richness and density. Both precipitation and temperature were positively correlated with soil total N, and total N was negatively correlated with vegetation richness. Both precipitation and temperature were negatively correlated with soil pH, and soil pH was negatively correlated with vegetation richness, but positively correlated with seed bank richness and density. Increasing precipitation and temperature would decrease seed bank diversity through direct effects as well as indirectly by decreasing vegetation diversity. Soil pH and total N emerged as the most important soil abiotic factors for seed bank diversity. Increasing precipitation and temperature under climate change may increase the extinction risk of some species in the seed bank by altering bet-hedging and risk-spreading strategies, which will degrade natural restoration ability and ultimately ecosystem resilience. This research is important because it identifies the potential underlying mechanistic basis of climate change impacts on seed banks through effects of aboveground vegetation and belowground biotic and abiotic factors.