Alpine Shrubland Research Articles

Characteristics and Driving Factors of Energy Balance over Different Underlying Surfaces in the Qinghai Plateau

The study of the surface energy balance characteristics of different ecosystems in the Qinghai Plateau is of great significance for a deeper understanding of land surface processes, the water cycle, and global climate change. This study aims to compare the seasonal variations in energy balance and partitioning of four typical ecosystems on the Qinghai Plateau—swamp meadows, subalpine mountain meadows, alpine shrublands, and alpine deserts. Mantel analysis and path analysis were used to explore the regulatory mechanisms of meteorological elements on energy fluxes and the Bowen ratio (β). The results showed the following: (1) Net radiation (Rn), sensible heat flux (H), and latent heat flux (LE) all exhibited a single-peak pattern of change, and the energy partitioning was closely related to the hydrothermal conditions. Swamp meadows and subalpine mountain meadows were dominated by LE throughout the year and the growing season, while H dominated in the non-growing season. Meanwhile, alpine shrublands and alpine deserts were dominated by H throughout the year. (2) β reflected the characteristics of turbulent fluxes variations and the moisture level of the underlying surface. Swamp meadows and subalpine mountain meadows were relatively moist, with the value of β all being less than 1. Alpine shrublands and deserts were comparatively arid, with the values of β all exceeding 1. The energy closure rate ranged from 48% to 90%, with better energy closure conditions observed during the growing season compared to the non-growing season. (3) Meteorological factors collectively regulated the variations in energy fluxes and its partitioning, with H and LE being primarily influenced by Rn, relative humidity (RH), and soil moisture (Ms). β was significantly affected by RH, Ms, and the saturated vapor pressure deficit (VPD). The sensitivity of the ecosystems to changes in fluxes increased with decreasing moisture, especially in alpine deserts, with Ms, VPD and RH being the most affected. Swamp meadows were significantly associated with air temperature (Ta), soil temperature (Ts), and wind speed; subalpine mountain meadows with Ta and Ts; and alpine shrublands with Ta. These results provided a basis for further analyses of the energy balance characteristics and partitioning differences of different ecosystems on the Qinghai Plateau.

Nitrogen addition reduces litter decomposition but does not affect litter production and chemistry in an alpine shrubland

Nitrogen addition reduces litter decomposition but does not affect litter production and chemistry in an alpine shrubland

The energy-limited water loss of an alpine shrubland on the northeastern Qinghai-Tibetan Plateau, China

Study regionAlpine shrubland on the northeastern Qinghai-Tibetan Plateau. Study focusWater provision ability is a pivotal ecological service of high-altitude alpine regions and is controlled by precipitation, evapotranspiration (ET), and soil water storage whereas the underlying ecohydrological processes remain highly unquantified. Here, we investigated continuous 19-year flux measurements to quantify the temporal patterns of ET and water budget (precipitation minus ET, P−ET), as well as 0-20 cm soil water storage change (ΔSWS). New hydrological insights for the regionAt a monthly scale, ET peaked in July (96.7 ± 26.4 mm, Mean ± S.D.) and averaged 41.7 ± 31.9 mm, whose variations were determined by the slope of the saturation vapor pressure curve at air temperature, air and soil temperatures, regardless of vegetation growth stage. P−ET averaged 18.3 ± 26.3 mm in August and September while stayed deficit during the other months. The variations in P−ET were controlled by precipitation in the May-October growing season whereas by ET in the non-growing season from November to April. ΔSWS peaked in May (28.8 ± 11.2 mm) and September (3.0 ± 2.7 mm) and almost accumulated to zero over the whole season. At annual scales, none of ET, P−ET, and ΔSWS changed significantly. ET averaged 512.2 ± 68.4 mm and exceeded precipitation (459.1 ± 58.4 mm), likely due to the lateral flow supply of uphill locations. The variations in ET were regulated directly by bulk canopy resistance and indirectly by net radiation. P−ET averaged −53.2 ± 95.4 mm and demonstrated a clear water deficit (−51.6 ± 21.0 mm) during the non-growing season. The variations of P−ET were driven jointly by precipitation and ET, with opposite but equivalent effects. The dominance of thermal conditions and energy availability on ET variability manifested an energy-limited feature of water loss in the alpine shrubland. The temporal patterns in P−ET elucidated that the alpine shrubland plays the water retention rather than water provision function through transforming variable precipitation input into stable ET loss.

Seasonal freeze‒thaw processes impact microbial communities of soil aggregates associated with soil pores on the Qinghai–Tibet Plateau

BackgroundSeasonal freeze‒thaw (FT) processes alter soil formation and cause changes in soil microbial communities, which regulate the decomposition of organic matter in alpine ecosystems. Soil aggregates are basic structural units and play a critical role in microbial habitation. However, the impact of seasonal FT processes on the distribution of microbial communities associated with soil pores in different aggregate fractions under climate change has been overlooked. In this study, we sampled soil aggregates from two typical alpine ecosystems (alpine meadow and alpine shrubland) during the seasonal FT processes (UFP: unstable freezing period, SFP: stable frozen period, UTP: unstable thawing period and STP: stable thawed period). The phospholipid fatty acid (PLFA) method was used to determine the biomass of living microbes in different aggregate fractions.ResultsThe microbial biomass of 0.25–2 mm and 0.053–0.25 mm aggregates did not change significantly during the seasonal FT process while the microbial biomass of > 2 mm aggregates presented a significant difference between the STP and UTP. Bacterial communities dominated the microbes in aggregates, accounting for over 80% of the total PLFAs. The microbial communities of soil aggregates in the surface layer were more sensitive to the seasonal FT process than those in other soil layers. In the thawing period, Gram positive bacteria (GP) was more dominant. In the freezing period, the ratio of Gram-positive to Gram-negative bacterial PLFAs (GP/GN) was low because the enrichment of plant litter facilitated the formation of organic matter. In the freezing process, pores of 30–80 μm (mesopores) favored the habitation of fungal and actinobacterial communities while total PLFAs and bacterial PLFAs were negatively correlated with mesopores in the thawing process.ConclusionsThe freezing process caused a greater variability in microbial biomass of different aggregate fractions. The thawing process increased the differences in microbial biomass among soil horizons. Mesopores of aggregates supported the habitation of actinobacterial and fungal communities while they were not conducive to bacterial growth. These findings provide a further comprehension of biodiversity and accurate estimation of global carbon cycle.

Differences and drivers of leaf stable carbon and nitrogen isotope in herbs under different vegetation types on the eastern Qinghai-Tibet Plateau

The natural abundance of stable carbon and nitrogen isotopes (δ13C and δ15N) in leaves can provide comprehensive information on the physiological and ecological processes of plants and has been widely used in ecological research. However, recent studies on leaf δ13C and δ15N have focused mainly on woody species, few studies have been conducted on herbs in different vegetation types, and their differences and driving factors are still unclear. In this study, we focused on the herbs in subalpine coniferous forests, alpine shrublands, and alpine mea-dows on the eastern Qinghai-Tibet Plateau, and investigated the differences in leaf δ13C and δ15N of herbs and the driving factors. The results showed that there were significant differences in leaf δ13C and δ15N values of herbs among different vegetation types, with the highest δ13C and δ15N values in alpine meadows, followed by alpine shrublands, and the lowest in subalpine coniferous forests. Using variation partitioning analysis, we revealed that differences in leaf δ13C and δ15N of herbs among various vegetation types were driven by both leaf functional traits and climate factors, with the contribution of leaf functional traits being relatively higher than that of climate factors. Hierarchical partitioning results indicated that mean annual temperature (MAT), chlorophyll content index, leaf nitrogen content per unit area (Narea), and leaf mass per area were the main drivers of leaf δ13C variations of herbs across different vegetation types, while the relative importance of Narea and MAT for variation in leaf δ15N of herbs was much higher than those other variables. There was a strong coupling relationship between leaf δ13C and δ15N as indicated by the result of the ordinary least squares regression. Our findings could provide new insights into understanding the key drivers of leaf δ13C and δ15N variations in herbs across different vegetation types.

Controls on use efficiency of plant nutrients along subtropical to alpine gradients on the Tibetan Plateau

AbstractQuestionKnowledge of how nutrient use strategies differ between forest trees and alpine shrubs/grasses is important to understand the mechanisms of vegetation changes from montane forests to alpine shrubs/grasslands along altitudinal gradients. We tested the hypothesis that, to maximize the nitrogen use efficiency (NUE) of canopy production, forest trees tend to have a higher mean residence time (MRT) of nitrogen in the plants through increased leaf life span, whereas alpine shrublands and grasslands tend to have higher nitrogen productivity through increased below‐ground biomass fraction. We further tested whether similar patterns are found in phosphorus use efficiency (PUE).LocationTwenty‐one sampling sites along Tibetan Alpine Vegetation Transects (TAVT) at altitudes from 1900 m to 4900 m.MethodsWe measured the maximum biomass of new canopy leaves and twigs and the concentrations of their nutrients N and P and associated ecosystem variables along the TAVT. NUE (PUE) was calculated as the product of nutrient productivity (dry matter production per unit N or P in new canopy leaves and twigs) and MRT (the ratio of foliage nutrient pool to annual nutrient uptake).ResultsWith increasing altitude, leaf life span increased in forest trees but decreased in shrublands and grasslands, while below‐ground fraction increased when vegetation changed from montane forests to alpine shrubs/grasslands. In forest trees, higher N‐MRT and P‐MRT and lower P productivity were associated with longer leaf life span and lower below‐ground fraction, while N productivity varied little. In alpine shrublands and grasslands, N‐MRT, P‐MRT and P productivity varied little with leaf life span and below‐ground fraction, while N productivity was positively correlated with below‐ground fraction.ConclusionsOur data supported the hypothesis, suggesting that NUE of canopy production would be a measure of changes in ecosystem functioning from montane forests to alpine shrublands and grasslands along altitudinal gradients. The findings provide an insight into the linkage between biogeochemistry and phytogeographic processes across ecosystems.

Exploring NDVI change patterns across the Tibetan Plateau at the hillslope scale using geomorphons

ABSTRACTWithin the Tibetan Plateau (TP), grassland degradation is of great concern, and there is a need to better understand the spatial variability and landscape patterns in grassland degradation across the TP. This study explores potential patterns in grassland degradation, estimated using Landsat-derived normalized difference vegetation index (NDVI) growing season change products generated by a prior study. Shuttle Radar Topography Mission (SRTM) digital terrain model (DTM)-derived geomorphons, topographic slope and the topographic position index (TPI) were used to explore grassland degradation and assess whether changes in growing season NDVI are associated with specific grasslands or landforms. An increase in NDVI within the TP was observed especially during the 1990 to 2018 and 2000 to 2018 time periods. Higher growing season median NDVI change values were found for the Southeast Tibet Shrublands and Meadows (SETSM) and TP Alpine Shrublands and Meadows (TPASM) grasslands in comparison to the other grassland types analysed, suggesting a more pronounced greening trend in the eastern portion of the plateau in comparison to the western portion. Small differences in NDVI change were observed for different geomorphon-based landforms, while topographic slope and TPI showed only modest correlations with NDVI change for all time periods and grassland types. More localized patterns of grassland degradation were masked by the widespread greening trends. Further analysis of only pixels that experienced a decreasing NDVI magnitude of less than or equal to −0.1 also generally suggested more change in the SETSM and TPASM. While this study generally supports the use of geomorphons as an analysis and aggregating unit for studying change patterns at the hillslope scale, more research is required to fully grasp the dynamics of landscape change at the hillslope scale in the TP.

Classification of forest and shrubland vegetation in central and eastern Euxine Turkey and SW Georgia

AbstractQuestionsWhat are the main vegetation types of forest and shrubland vegetation in central and eastern Euxine Turkey and SW Georgia? What are the main environmental factors affecting their diversity? What is their syntaxonomic position? Can we integrate them into the European vegetation classification system?LocationCentral and eastern Euxine Turkey and SW Georgia.MethodsWe collected 3104 vegetation plots of forest and shrubland vegetation in the study region and performed Two‐Way Indicator Species Analysis (TWINSPAN) classification. We described vegetation types based on the classification results, expert knowledge and information from literature sources. We defined diagnostic species and prepared distribution maps for each vegetation type. To determine the most significant environmental variables on floristic differentiation, we used canonical correspondence analysis. Detrended correspondence analysis with passive projection of most significant environmental variables was run to interpret the environmental variation of vegetation types.ResultsThe studied vegetation was divided into 29 vegetation types related to seven main vegetation groups: relict Mediterranean forests and shrubland (mainly along the coastline, beside some inland localities), lowland to submontane forests, central Euxine mountain forests, eastern Euxine (Colchic) mountain forests, subeuxine forests, azonal riparian forests and subalpine and alpine shrubland. Elevation is the most important factor causing the differentiation in vegetation. It is followed by longitude and latitude. Among climatic variables, temperature seasonality, annual precipitation and precipitation of the wettest quarter are the most significant factors for vegetation differentiation. These factors correlate with the reduction of maritime climate and geomorphological features.ConclusionsVegetation types mostly correspond to the syntaxa accepted in the EuroVegChecklist. However, some of them do not appear in the EuroVegChecklist since they appear only beyond Europe. We described three syntaxa as new: Abietion equi‐trojani, Querco cerridis‐Carpinion orientalis and Piceo orientalis‐Fagenion orientalis. The study revealed high vegetation diversity of the region that should be taken into consideration in ecosystem management and used as a reference in restoration and mitigation of the effects of global changes.

Divergent environmental responses of long-term variations in evapotranspiration over four grassland ecosystems in China based on eddy-covariance measurements

Understanding the long-term variation in evapotranspiration (ET) for the spatially distributed grasslands is crucial for the accurate prediction of ET response to climate change. In this study, we analyzed the interannual variability (IAV) of ET and its responses to environmental conditions at four grassland ecosystems across a wide range of climatic and biome conditions based on the long-term (9–11 years) eddy-covariance measurements. The four ecosystems encompassed the most prevalent grassland vegetations in China, containing a typical temperate steppe, an alpine meadow-steppe, an alpine shrubland meadow, and an alpine marsh meadow. The IAVs of annual ET at the typical temperate steppe and the alpine meadow-steppe were primarily affected by either change in precipitation (P) or relative humidity (RH). Leaf area index (LAI) was the dominant factor controlling the IAVs of annual and growing-season ET at the alpine shrubland meadow, and the IAV of LAI was significantly correlated with P variation. As to the alpine marsh meadow, net radiation turned to be the dominant factor for the IAV of annual ET, additionally with significant effects from water supply condition (P and RH) on the IAV of growing-season ET. Similar environmental responses were also found for the IAVs of mean surface conductance (gs) across the sites. Specifically, annual and growing-season mean gs significantly increased with increases in LAI at the alpine shrubland meadow, and appeared to be more sensitive to changes in water availability and VPD at the typical temperate steppe and the alpine meadow-steppe. Significant linear relationships were also observed among the IAVs of mean Priestley-Taylor coefficient (α = ET/ETeq, where ETeq is the equilibrium evaporation), decoupling coefficient (Ω), and gs on both the annual and growing-season basis in this study. Moreover, the variabilities of annual mean gs, Ω, and α further demonstrated the energy-limited conditions at the alpine marsh meadow, and the overall water-limited conditions at the other three grasslands. This study reveals the divergent environmental responses of long-term ET variations over grassland ecosystems, and contributes to the comprehensive understanding on the ET process and modeling efforts as well.

Different responses of soil element contents and their stoichiometry (C: N: P) to different grazing intensity on the Tibetan Plateau shrublands

Potentilla fruticosa, a major alpine shrubland type, is widely distributed across the Tibetan Plateau, and grazing is the most common disturbance in the shrublands of P. fruticosa. However, soil organic carbon (SOC), soil total nitrogen (STN), soil total phosphorus (STP), and their stoichiometry under different grazing intensities were unclear. In our study, we explored SOC, STN, STP, their stoichiometry, and their controlling factors in the grazing disturbance of heavy grazing (HG), moderate grazing (MG), light grazing (LG), and no grazing (NG) conditions in the Tibetan Plateau P. fruticosa shrublands. The grazing intensities were mainly assessed by considering the shrublands’ ground cover, the indicators of the road density, the distance between sampling sites and cowshed or sheep shed, the amounts of cow and sheep dung, and vegetation that had been gnawed and stampeded. Our results indicated that soil physical properties of soil temperature and bulk density have decreasing trends with decreasing grazing intensities from HG to NG. The SOC, STN, STP, and soil C:N and C:P ratios have increasing trends with decreasing grazing intensities from HG to NG, while the changes in soil N:P ratio were relatively stable along grazing intensities. Our results indicated that HG generally had stronger effects on SOC, STN, and soil C:N and C:P ratios than NG, indicating substantial effects of grazing disturbance on biogeochemical cycles of SOC and STN in the shrubland ecosystems. Therefore, for the alpine shrubland of P. fruticosa, the HG should be avoided for sustainable cycling of soil nutrients and the balance of soil nutrient stoichiometry. The grazing types can directly affect plant conditions, and plant conditions can directly affect soil physical and chemical properties and litter standing crops. Finally, soil physicochemical properties and litter standing crop resulting from different grazing intensities directly control SOC, STN, and STP. For the soil stoichiometry, the soil’s physical and chemical properties resulting from different grazing intensities have direct impacts on soil C:P and N:P ratios.

A dataset of the observations of carbon, water and heat fluxes over an alpine shrubland in Haibei (2011–2020)

Alpine shrubland is one of the important vegetation types on the Qinghai-Tibet Plateau, which mainly lies in the shady or semi-shady slope of snowpack mountains or the high-altitude alluvium and diluvium on plains. It plays a crucial role in carbon sequestration, water conservation and climate regulation. Since 2002, Haibei National Field Research Station for Alpine Grassland (Haibei Station) has been using eddy covariance techniques to continuously observe the carbon, water and heat exchange between an alpine Potentilla fruticosa shrubland ecosystem and the atmosphere and has accumulated nearly 20-year data. On the basis of the previous publication of relevant data from 2003 to 2010, the carbon, we further released water and heat fluxes of the alpine shrubland and supplementary meteorological data from 2011 to 2020. This dataset consists of the subsets of meteorological factors, covering air temperature, air relative humidity, water vapor pressure, wind speed, wind direction, ambient pressure, total solar radiation, net radiation, photosynthetically active radiation, precipitation, soil temperature, and soil moisture, as well as net ecosystem CO2 exchange, ecosystem respiration, gross ecosystem CO2 exchange, latent heat flux, and sensible heat flux. The temporal resolutions of the dataset include half-hourly, daily, monthly, and yearly scales. This dataset can not only be used to scientifically evaluate the environmental drivers and evolution trends of the ecological functions of carbon, water and heat in alpine shrub ecosystems, but also provide ground data support for parameter validation and optimization of remote sensing-based ecological process models.

Loss of growth resilience towards the alpine shrubline

The loss of resilience and prolonged recovery times after extreme climate events can be used as early warning signals of impending tipping points or abrupt irreversible changes. However, evidence of such critical slowing down in growth series is still lacking in terrestrial ecosystems under harsh environmental conditions such as alpine shrublands. Alpine shrublines, at which shrublands shift into grasslands under climate thresholds, constitute unique settings to look for critical slowing down phenomena. On the northeastern Tibetan Plateau, we sampled Salix oritrepha Schneid. (willow shrubs) in three elevational sites along each of two transects (QL, Qilian; HSX, Huashixia) running from alpine treeline position to shrubline. We then quantified the radial growth resilience and recovery indices of alpine willow shrubs to cold spells along elevational gradients by using dendrochronology. Willow growth was primarily constrained by low winter temperatures in QL and low summer temperatures in HSX. Cold spell events were detected in 2008 and 2011 in HSX and QL sites, respectively. Because temperature decreases with elevation, both growth resilience and recovery indices differed within each transect, showing a decreasing trend with elevation. Growth in 78% and 56% willow shrubs recovered back to the level before cold spell within two years in QL and HSX sites, respectively. Moreover, the average growth recovery period was longer at the shrubline (ca. 2–4 years) than near the treeline position (ca. 1–2 years). Critical slowing down theory could thus be applied to assess growth resilience towards the alpine shrubline, a thermal threshold for shrub growth. Our results and framework highlight the priority to directly measure the resilience metrics and also offer a promising perspective that the critical slowing down phenomenon may be universal in various ecosystems.

Relationships between modern pollen and vegetation and climate on the eastern Tibetan Plateau

The Tibetan Plateau ecosystem is fragile and sensitive to climate change. Understanding the relationships between modern pollen and the vegetation and climate of the region is critical for the evaluation of ecological processes. Here, we explore modern pollen assemblages of typical land-cover types at a large spatial scale by analyzing 36 surface samples from the southeastern Tibetan Plateau, supplemented by typical desert, desert-steppe, and steppe meadow transition data selected from the Chinese Surface Pollen Database, giving a total of 75 samples. We used redundancy analysis (RDA) to explore the responses of vegetation in the assemblages to regional climate. Our results show that pollen assemblages generally reflect the vegetation composition: assemblages from alpine meadow samples are dominated by Cyperaceae, Asteraceae, Rosaceae, and Polygonaceae; alpine shrublands mainly comprise Fabaceae, Rosaceae, Ericaceae, and Quercus (Q. spinosa); and coniferous forest surface samples mainly comprise Picea, Abies, Pinus, and Betulaceae. Our RDA shows that mean annual precipitation (MAP) is the main meteorological factor affecting the pollen assemblage and vegetation type; MAP positively correlates with percentages of Cyperaceae, Poaceae, Rosaceae, and Asteraceae, and negatively correlates with percentages of Chenopodiaceae, Ephedraceae, Nitraria, and Tamaricaceae. The ratio of Artemisia to Chenopodiaceae is a useful indicator to distinguish temperate desert from other land-cover types on the Tibetan Plateau, while the ratio of Cyperaceae + Asteraceae to Artemisia + Chenopodiaceae can be used to distinguish arid desert from other land-cover types, and may provide a useful altitude index for the eastern Tibetan Plateau.

Contrasting responses of soil microbial biomass and extracellular enzyme activity along an elevation gradient on the eastern Qinghai-Tibetan Plateau.

Soil microbial community composition and extracellular enzyme activity are two main drivers of biogeochemical cycling. Knowledge about their elevational patterns is of great importance for predicting ecosystem functioning in response to climate change. Nevertheless, there is no consensus on how soil microbial community composition and extracellular enzyme activity vary with elevation, and little is known about their elevational variations on the eastern Qinghai-Tibetan Plateau, a region sensitive to global change. We therefore investigated the soil microbial community composition using phospholipid fatty acids (PLFAs) analysis, and enzyme activities at 2,820 m (coniferous and broadleaved mixed forest), 3,160 m (dark coniferous forest), 3,420 m (alpine dwarf forest), and 4,280 m (alpine shrubland) above sea level. Our results showed that soil microbial community composition and extracellular enzyme activities changed significantly along the elevational gradient. Biomass of total microbes, bacteria, and arbuscular mycorrhizal fungi at the highest elevation were the significantly lowest among the four elevations. In contrast, extracellular enzyme activities involved in carbon (C)-, nitrogen (N)-, and phosphorus (P)- acquiring exhibited the maximum values at the highest elevation. Total nutrients and available nutrients, especially P availability jointly explained the elevational pattern of soil microbial community, while the elevational variation of extracellular enzyme activities was dependent on total nutrients. Microbial metabolism was mainly C- and P-limited with an increasing C limitation but a decreasing P limitation along the elevational gradient, which was related significantly to mean annual temperature and total P. These results indicated a vital role of soil P in driving the elevational patterns of soil microbial community and metabolism. Overall, the study highlighted the contrasting responses of soil microbial biomass and extracellular enzyme activities to elevation, possibly suggesting the differences in adaption strategy between population growth and resource acquisition responding to elevation. The results provide essential information for understanding and predicting the response of belowground community and function to climate change on the eastern Qinghai-Tibetan Plateau.

Effects of warming on the stoichiometry of soil microbial biomass and extracellular enzymes in an alpine shrubland

The alpine shrublands on the Qinghai-Tibetan Plateau are experiencing significant warming. Revealing the effects of warming on the soil microbial metabolism will provide insights into the dynamics of soil carbon (C) and nutrient cycling in these alpine ecosystems. A five-year warming experiment (+1.3 °C, using open-top chambers) was conducted in a Sibiraea angustata-dominated alpine shrubland on the eastern Qinghai-Tibetan Plateau. We examined the effects of warming on the stoichiometry of topsoil (0–15 cm) microbial biomass, extracellular enzyme activity and their relationships with soil physicochemical properties. The results showed that the effects of warming on the soil microbial biomass C, nitrogen (N) and phosphorus (P) varied with the growing season. Warming increased the activity levels of C-acquiring enzymes (β-glucosidase, BG) by 7.5 %-15.1 % throughout the growing season (P < 0.01) and the activity of polyphenol oxidase (PPO) by 6.5 % during the early growing season (P < 0.05), indicating that warming would strengthen microbial C limitation and increase degradation of soil labile and recalcitrant C components. Warming increased the activity of P-acquiring enzyme (acid phosphatase, AP) by 14.6 % (P < 0.01) and triggered microbial P limitation during the late growing season, which might be a result of an increasing alpine shrub P uptake. Warming did not affect the total activity levels of N-acquiring enzymes throughout the growing season (P > 0.05), mainly due to higher soil N availability. Warming did not affect the stoichiometry of soil extracellular enzymes throughout the growing season (P > 0.05), except for a decreasing in the enzyme N:P ratio during the early growing season (P < 0.05). Moreover, the microclimate (air and soil temperature), available soil nutrients (soil dissolved organic C and available P) and soil pH accounted for most of the variation in soil microbial biomass, extracellular enzyme activity and their stoichiometry in response to warming. Our results suggest that warming may trigger and strengthen soil microbial C and P limitation during the growing season, accelerating soil C loss and nutrient cycling in alpine shrublands on the Qinghai-Tibetan Plateau.

Contrasting responses of microbial diversity and community structure in decaying root bark and xylem to N addition in an alpine shrubland

Microbial degradation of root components (root bark and xylem) is a vital biogeochemical process that may be influenced by N deposition because N often limits microbial growth. However, there is a lack of information on microbial diversity and community structure during long-term decomposition of root bark and xylem under N addition, which seriously hinders our understanding of microbial decomposition of the root. In this study, litterbags containing roots from a dominant alpine shrub species (Sibiraea angustata) were buried in soil and sampled after 8 years of decomposition and four levels of N addition. The bacterial and fungal diversity significantly differed between root bark and xylem, due to their distinct C chemistry and water content. The microbial richness in the root bark decreased linearly with increasing N levels, whereas the opposite trend was observed in the root xylem. Additionally, high levels of N significantly altered the microbial community structure in the root bark but not in the root xylem. Despite these variations in overall microbial community patterns, N addition did not affect the root-decomposition rate and corresponding recalcitrant C in the root bark and xylem, likely due to the unchanged relative abundances of key functional guilds involved in root decomposition (i.e. litter saprotrophs, cellulolytic bacteria, and ligninolytic bacteria). Given the contrasting microbial responses to N addition in root components, our results suggest that the theory on root decomposition should consider different microbial controls in the decomposition of root bark and xylem.

Long-term warming reduces surface soil organic carbon by reducing mineral-associated carbon rather than “free” particulate carbon

Soils in alpine ecosystems store a large amount of carbon. However, the responses of soil organic carbon (SOC) to long-term warming and the microbial mechanisms underlying SOC dynamics remain poorly understood. Here, we investigated the effects of 20-year warming (1997–2017) on SOC pools in an alpine shrubland on the Qinghai-Tibetan Plateau. We analyzed the plant and microbial communities, as well as soil chemistry, particularly the SOC of bulk soil and its two physical fractions. Warming did not alter plant biomass (either above- or below-ground). However, warming decreased overall microbial biomass carbon and nitrogen, the biomass of fungi (indicated by phospholipid fatty acids, amino sugars, and OTU richness), and potential activities of hydrolytic enzymes (especially β-1,4-glucosidase), in the surface 0–10 cm soil. Moreover, warming reduced the SOC in bulk soil (by 21%) and the heavy fraction (by 25%), but did not change the SOC in the light fraction. Importantly, there were strong positive correlations between the SOC in bulk soil and in the heavy fraction with the total microbial and fungal necromass, suggesting that the warming-induced loss of soil carbon was mainly due to the suppression of the microbial pathway of soil carbon formation. In deeper layers (10–50 cm) of the shrubland, however, warming (to a much lower degree than that in the surface soil) had only a minor effect on soil carbon pools and the microbial community. A global meta-analysis further showed that warming on average did not significantly change the SOC in different fractions (bulk SOC, active C, and stable C) in the surface soil. Taken together, these results show that long-term warming leads to significant loss of surface soil carbon (mostly in the mineral-associated heavy fraction), likely due to suppressing the microbial (particularly fungal) pathway to soil carbon formation and/or stimulating the utilization of stable soil carbon (i.e. heavy fraction) by microbes in the alpine shrubland.

Increased plant growth may offset soil carbon loss caused by warming in an alpine Sibiraea angustata shrub ecosystem on the eastern Qinghai-Tibet Plateau

The alpine shrub ecosystem on the Qinghai-Tibet Plateau, which reserves large amounts of carbon (C) and functions as an important potential C sink, is experiencing significant warming. However, we still lack a thorough understanding of the effects of warming on ecosystem C storage and allocation in above- and belowground components of plants and soils in these alpine shrub ecosystems. In this study, we employed a four-year field warming experiment in an alpine shrubland dominated by Sibiraea angustata on the eastern Qinghai-Tibetan Plateau. We investigated the effects of warming on C storage and allocation among the above- (i.e., leaf and stem) and belowground parts (i.e., coarse and fine roots) of shrubs and the above- and belowground parts of grasses and soils. Our results showed that warming significantly increased shrub leaf C storage by 18.8%, but shrub stem C storage did not respond to warming. Warming significantly increased the C storage of shrub coarse and fine roots by 7.7% and 139.4%, respectively. Warming significantly increased the C storage of the above- and belowground parts of grasses by 60.4% and 130.5%, respectively. As a result, warming significantly increased the C storage of both the above- and belowground parts of total plants by 8.8% and 49.1%, respectively. Moreover, warming significantly increased the C root:shoot ratios of alpine shrubs and grasses. Taken together, warming significantly increased the C storage of total plants by 1.23 kg C m−2, which primarily resulted from the belowground parts of plants (1.07 kg C m−2), and warming-induced increases in C storage of alpine shrub fine roots (0.86 kg C m−2) could approximately offset soil C losses (0.99 kg C m−2), as indicated by decreasing soil C storage under warming. Therefore, warming increased the ecosystem C sink by 0.24 kg C m−2. Our results highlight that alpine shrub fine roots play a dominant role in regulating the dynamics of ecosystem C balance and cycling in response to warming. Future climate warming will stimulate ecosystem C sequestration, largely because of increasing the growth of alpine shrub fine roots in these alpine shrublands on the Qinghai-Tibet Plateau.

Vegetation Classification and Distribution Patterns in the South Slope of Yarlung Zangbo Grand Canyon National Nature Reserve, Eastern Himalayas

Yarlung Zangbo Grand Canyon National Nature Reserve has the most complete vertical vegetation belts in China. However, identification and distribution of vertical vegetation belts is still uncertain and in debate. To explore the above issues, 190 plots were surveyed within the reserve from 2019 to 2021. Based on the vegetation plot data, cluster analysis, ordination analysis, and biodiversity statistics were performed to reveal the structure of vertical vegetation belts–the driving factors of vegetation distribution–to describe the main biodiversity patterns. Five vertical vegetation belts were identified by clustering. NMDS ordination showed that the main factor of vegetation distribution is elevation. Based on the results of the analysis and previous literature, a new scheme of vertical vegetation belts in the south slope of the reserve was proposed. There was a lower montane seasonal rainforest belt (600–1100 m), a lower montane evergreen broadleaf forest belt (1100–1800 m), a middle montane semi-evergreen broadleaf forest belt (1800–2400 m), a subalpine evergreen needleleaf forest belt (2400–3800 m), a alpine shrubland and meadow belt (3800–4400 m), an alpine sparse vegetation belt (4400–4800 m), and a nival belt (4800–7782 m). Among them, the seasonal rainforest belts are the northernmost distribution of this type, and the semi-evergreen broadleaf forest belts exist only in the Eastern Himalayas. The study showed a unimodal pattern in plant species diversity, the peak of which is about 1900 m. The middle montane semi-evergreen broadleaf forest belt had the highest species diversity in the reserve. This study settled the issues regarding the vertical vegetation belts, the main drivers of vegetation and assessment of plant species diversity in the south slope of the Yarlung Zangbo Grand Canyon National Nature Reserve. It provides essential support for the management and conservation of these ecosystems in the reserve.

Distribution Characteristics and Controlling Factors of Soil Total Nitrogen: Phosphorus Ratio Across the Northeast Tibetan Plateau Shrublands.

Nitrogen (N) and phosphorus (P) stoichiometry have significant effects on nutrient cycles in terrestrial ecosystems. However, our understanding of the patterns and the driving factors of soil N:P ratios in the Tibetan Plateau shrublands remains limited. Our study aimed to quantify the distribution of soil N:P ratio and its controlling factors based on soil, plant, and climate factors from 59 sites in shrublands across the northeast Tibetan Plateau. The kriging interpolation method was used to quantify the soil N:P distribution. Spatially, the soil N:P ratio was higher in the south than in the north and lower in the west than in the east. The soil N:P ratio in the northeast Tibetan Plateau shrublands was mainly explained by edaphic factors, which also played an important role in regulating the effects of plant and climate factors on soil N:P ratios. Mean annual precipitation, instead of mean annual temperature, significantly controlled the soil N:P ratios, and its effect on the pattern of soil N:P ratios differed between alpine shrublands and desert shrublands. The N:P ratios of different organs in shrublands also played different roles in shaping the soil N:P ratios in alpine and desert shrublands. These results provide support for the hypothesis that edaphic factors were the dominant drivers of spatial variation in soil N:P ratios across the northeast Tibetan Plateau shrublands, and our study contributes to a deeper understanding of biogeochemical cycling at high altitudes.