Tianxiang Luo

Little change in the fir tree‐line position on the southeastern Tibetan Plateau after 200 years of warming

• As one of the worlds highest natural tree lines, the Smith fir (Abies georgei var. smithii) tree line on the southeastern Tibetan Plateau is expected to vary as a function of climate warming. However, the spatial patterns and dynamics of the Smith fir tree line are not yet well understood. • Three rectangular plots (30 m × 150 m) were established in the natural alpine tree-line ecotone on two north-facing (Plot N1, 4390 m asl; Plot N2, 4380 m asl) and one east-facing (Plot E1, 4360 m asl) slope. Dendroecological methods were used to monitor the tree-line patterns and dynamics over a 50-yr interval. • The three study plots showed a similar pattern of regeneration dynamics, characterized by increased recruitment after the 1950s and an abrupt increase in the 1970s. Smith fir recruitment was significantly positively correlated with both summer and winter temperatures. However, Smith fir tree lines do not show a significant upward movement, despite warming on the Tibetan Plateau. • The warming in the past 200 yr is already having a significant impact on the population density of the trees, but not on the position of the Smith fir tree line.

Leaf traits and associated ecosystem characteristics across subtropical and timberline forests in the Gongga Mountains, Eastern Tibetan Plateau

Knowledge of how leaf characteristics might be used to deduce information on ecosystem functioning and how this scaling task could be done is limited. In this study, we present field data for leaf lifespan, specific leaf area (SLA) and mass and area-based leaf nitrogen concentrations (Nmass, Narea) of dominant tree species and the associated stand foliage N-pool, leaf area index (LAI), root biomass, aboveground biomass, net primary productivity (NPP) and soil available-N content in six undisturbed forest plots along subtropical to timberline gradients on the eastern slope of the Gongga Mountains. We developed a methodology to calculate the whole-canopy mean leaf traits to include all tree species (groups) in each of the six plots through a series of weighted averages scaled up from leaf-level measurements. These defined whole-canopy mean leaf traits were equivalent to the traits of a leaf in regard to their interrelationships and altitudinal trends, but were more useful for large-scale pattern analysis of ecosystem structure and function. The whole-canopy mean leaf lifespan and leaf Nmass mainly showed significant relationships with stand foliage N-pool, NPP, LAI and root biomass. In general, as elevation increased, the whole-canopy mean leaf lifespan and leaf Narea and stand LAI and foliage N-pool increased to their maximum, whereas the whole-canopy mean SLA and leaf Nmass and stand NPP and root biomass decreased from their maximum. The whole-canopy mean leaf lifespan and stand foliage N-pool both converged towards threshold-like logistic relationships with annual mean temperature and soil available-N variables. Our results are further supported by additional literature data in the Americas and eastern China.

Spatial patterns of Smith fir alpine treelines on the south-eastern Tibetan Plateau support that contingent local conditions drive recent treeline patterns

Background: Recent work has shown little change in the position of the Smith fir treeline on the south-eastern Tibetan Plateau in response to global warming. However, the relationship between tree distribution patterns within the treeline ecotone and low responsiveness is unknown, and additional constraints than climate might be major drivers of these patterns (e.g. microsite availability for regeneration). Aims: To characterise the spatial patterns of Smith fir alpine treelines and to infer the underlying processes driving their dynamics. Methods: We investigated spatial patterns of Smith fir trees across two treeline ecotones in the Sygera Mountains, south-eastern Tibetan Plateau. The O(r)-ring statistic was used to analyse the univariate and bivariate spatial point patterns of three size classes (adults, juveniles and seedlings). Results: Mature trees presented random spatial patterns. Clusters of juveniles and seedlings colonised areas not occupied by mature trees. Seedlings were clustered and established preferentially near juvenile firs, Rhododendron mats and over moss–lichen and organic matter substrates, indicating the importance of microsite availability for successful Smith fir recruitment. Conclusions: Local factors such as microsite availability may play a major role in driving recent Smith fir treeline patterns and determine the lack of significant warming-induced upward shifts of these ecotones.

Linking carbon supply to root cell-wall chemistry and mechanics at high altitudes in Abies georgei

BACKGROUND AND AIMS The mobile carbon supply to different compartments of a tree is affected by climate, but its impact on cell-wall chemistry and mechanics remains unknown. To understand better the variability in root growth and biomechanics in mountain forests subjected to substrate mass movement, we investigated root chemical and mechanical properties of mature Abies georgei var. smithii (Smith fir) growing at different elevations on the Tibet-Qinghai Plateau. METHODS Thin and fine roots (0·1-4·0 mm in diameter) were sampled at three different elevations (3480, 3900 and 4330 m, the last corresponding to the treeline). Tensile resistance of roots of different diameter classes was measured along with holocellulose and non-structural carbon (NSC) content. KEY RESULTS The mean force necessary to break roots in tension decreased significantly with increasing altitude and was attributed to a decrease in holocellulose content. Holocellulose was significantly lower in roots at the treeline (29·5 ± 1·3 %) compared with those at 3480 m (39·1 ± 1·0 %). Roots also differed significantly in NSC, with 35·6 ± 4·1 mg g(-1) dry mass of mean total soluble sugars in roots at 3480 m and 18·8 ± 2·1 mg g(-1) dry mass in roots at the treeline. CONCLUSIONS Root mechanical resistance, holocellulose and NSC content all decreased with increasing altitude. Holocellulose is made up principally of cellulose, the biosynthesis of which depends largely on NSC supply. Plants synthesize cellulose when conditions are optimal and NSC is not limiting. Thus, cellulose synthesis in the thin and fine roots measured in our study is probably not a priority in mature trees growing at very high altitudes, where climatic factors will be limiting for growth. Root NSC stocks at the treeline may be depleted through over-demand for carbon supply due to increased fine root production or winter root growth.

Leaf unfolding of Tibetan alpine meadows captures the arrival of monsoon rainfall

The alpine meadow on the Tibetan Plateau is the highest and largest pasture in the world, and its formation and distribution are mainly controlled by Indian summer monsoon effects. However, little is known about how monsoon-related cues may trigger spring phenology of the vast alpine vegetation. Based on the 7-year observations with fenced and transplanted experiments across lower to upper limits of Kobresia meadows in the central plateau (4400–5200 m), we found that leaf unfolding dates of dominant sedge and grass species synchronized with monsoon onset, regardless of air temperature. We also found similar patterns in a 22-year data set from the northeast plateau. In the monsoon-related cues for leaf unfolding, the arrival of monsoon rainfall is crucial, while seasonal air temperatures are already continuously above 0 °C. In contrast, the early-emerging cushion species generally leafed out earlier in warmer years regardless of precipitation. Our data provide evidence that leaf unfolding of dominant species in the alpine meadows senses the arrival of monsoon-season rainfall. These findings also provide a basis for interpreting the spatially variable greening responses to warming detected in the world’s highest pasture, and suggest a phenological strategy for avoiding damages of pre-monsoon drought and frost to alpine plants.

Low rainfall‐induced shift in leaf trait relationship within species along a semi‐arid sandy land transect in northern China

It is unclear whether the shift in leaf traits between species of high- and low-rainfall sites is caused by low rainfall or by species replacement, because leaf traits vary substantially among species and sites. Our objective was to test if the within-species relationship between specific leaf area (SLA) and leaf N concentration (N(mass) ) shifts across a rainfall gradient in the semi-arid sandy lands of northern China. Data for SLA and N(mass) of dominant species and related canopy and soil variables were collected from 33 plots along a rainfall transect (270-390 mm) having similar temperatures in the Mu Us, Inner Mongolia. We further investigated the generality of Mu Us data using 12 additional plots in the southeastern Qaidam Basin, Qinghai. Artemisia ordosica is a widespread species in both regions. Across and within species, the positive SLA-N(mass) relationship shifted between two plant groups in the lowest rainfall plots (270 mm) and other higher rainfall plots (320-390 mm), which was confirmed by additional data from Qinghai. For A. ordosica populations, leaf area index (LAI) decreased and N(mass) increased with decreasing rainfall, while the foliage N pool and SLA varied little. Rainfall was the limiting factor that determined variations in N(mass) and LAI. Accordingly, N(mass) /SLA ratios continually increased with decreasing LAI along the rainfall gradient (r = -0.76, P < 0.001). Results indicate a low rainfall-induced shift in the SLA-N(mass) relationship associated with changes in LAI and foliage N pool, suggesting a link between leaf characteristics and ecosystem function.

Seasonal variations in leaf δ13C and nitrogen associated with foliage turnover and carbon gain for a wet subalpine fir forest in the Gongga Mountains, eastern Tibetan Plateau

It is still unclear to what extent variations in foliar δ13C and nitrogen can be used to detect seasonal changes in canopy productivity. We hypothesize that in a wet and cloudy fir forest, seasonally higher litterfall and lower leaf area index (LAI) are correlated with higher mass-based leaf nitrogen (Nmass) and net primary productivity (NPP), while foliar δ13C may change with specific leaf area (SLA), area-based leaf nitrogen (Narea), and/or starch concentration. In order to test our hypotheses, stand-level litterfall and the means of δ13C, Nmass, Narea, SLA, and starch concentration of canopy needles for a wet and cloudy Abies fabri forest in the Gongga Mountains were monthly measured during the growing season. Seasonal estimates of LAI were obtained from our previous work. A conceptual model was used to predict seasonal NPP of the fir forest. Seasonal mean δ13C and Nmass and climatic variables were used as inputs. The δ13C across 1–7-year-old needles increased from May to September associated with decreasing SLA and increasing Narea. There were no significant differences in seasonal starch concentration. With increasing litterfall and decreasing LAI, seasonal mean Nmass increased, while the δ13C varied little. The simulated NPP increased with increasing litterfall and related traits of Nmass and Narea. Our data generally supported the hypotheses. The results also suggest that in the forest with relatively moist and cloudy environment, the largest fraction of annual carbon gain may occur in the early part of the growing season when higher litterfall results in higher Nmass of canopy leaves.

Seasonal Shift in Climatic Limiting Factors on Tree Transpiration: Evidence from Sap Flow Observations at Alpine Treelines in Southeast Tibet

Alpine and northern treelines are primarily controlled by low temperatures. However, little is known about the impact of low soil temperature on tree transpiration at treelines. We aim to test the hypothesis that in cold-limited forests, the main limiting factors for tree transpiration switch from low soil temperature before summer solstice to atmospheric evaporative demand after summer solstice, which generally results in low transpiration in the early growing season. Sap flow, meteorological factors and predawn needle water potential were continuously monitored throughout one growing season across Smith fir (Abies georgei var. smithii) and juniper (Juniperus saltuaria) treelines in southeast Tibet. Sap flow started in early May and corresponded to a threshold mean air-temperature of 0°C. Across tree species, transpiration was mainly limited by low soil temperature prior to the summer solstice but by vapor pressure deficit and solar radiation post-summer solstice, which was further confirmed on a daily scale. As a result, tree transpiration for both tree species was significantly reduced in the pre-summer solstice period as compared to post-summer solstice, resulting in a lower predawn needle water potential for Smith fir trees in the early growing season. Our data supported the hypothesis, suggesting that tree transpiration mainly responds to soil temperature variations in the early growing season. The results are important for understanding the hydrological response of cold-limited forest ecosystems to climate change.

Optimal balance of water use efficiency and leaf construction cost with a link to the drought threshold of the desert steppe ecotone in northern China

BACKGROUND AND AIMS In arid environments, a high nitrogen content per leaf area (Narea) induced by drought can enhance water use efficiency (WUE) of photosynthesis, but may also lead to high leaf construction cost (CC). Our aim was to investigate how maximizing Narea could balance WUE and CC in an arid-adapted, widespread species along a rainfall gradient, and how such a process may be related to the drought threshold of the desert-steppe ecotone in northern China. METHODS Along rainfall gradients with a moisture index (MI) of 0·17-0·41 in northern China and the northern Tibetan Plateau, we measured leaf traits and stand variables including specific leaf area (SLA), nitrogen content relative to leaf mass and area (Nmass, Narea) and construction cost (CCmass, CCarea), δ(13)C (indicator of WUE), leaf area index (LAI) and foliage N-pool across populations of Artemisia ordosica KEY RESULTS In samples from northern China, a continuous increase of Narea with decreasing MI was achieved by a higher Nmass and constant SLA (reduced LAI and constant N-pool) in high-rainfall areas (MI > 0·29), but by a lower SLA and Nmass (reduced LAI and N-pool) in low-rainfall areas (MI ≤ 0·29). While δ(13)C, CCmass and CCarea continuously increased with decreasing MI, the low-rainfall group had higher Narea and δ(13)C at a given CCarea, compared with the high-rainfall group. Similar patterns were also found in additional data for the same species in the northern Tibetan Plateau. The observed drought threshold where MI = 0·29 corresponded well to the zonal boundary between typical and desert steppes in northern China. CONCLUSIONS Our data indicated that below a climatic drought threshold, drought-resistant plants tend to maximize their intrinsic WUE through increased Narea at a given CCarea, which suggests a linkage between leaf functional traits and arid vegetation zonation.

Contrasting changes in above- and below-ground biomass allocation across treeline ecotones in southeast Tibet

Under conditions of a warmer climate, the advance of the alpine treeline into alpine tundra has implications for carbon dynamics in mountain ecosystems. However, the above- and below-ground live biomass allocations among different vegetation types within the treeline ecotones are not well investigated. To determine the altitudinal patterns of above-/below-ground carbon allocation, we measured the root biomass and estimated the above-ground biomass (AGB) in a subalpine forest, treeline forest, alpine shrub, and alpine grassland along two elevational transects towards the alpine tundra in southeast Tibet. The AGB strongly declined with increasing elevation, which was associated with a decrease in the leaf area index and a consequent reduction in carbon gain. The fine root biomass (FRB) increased significantly more in the alpine shrub and grassland than in the treeline forest, whereas the coarse root biomass changed little with increasing altitudes, which led to a stable below-ground biomass (BGB) value across altitudes. Warm and infertile soil conditions might explain the large amount of FRB in alpine shrub and grassland. Consequently, the root to shoot biomass ratio increased sharply with altitude, which suggested a remarkable shift of biomass allocation to root systems near the alpine tundra. Our findings demonstrate contrasting changes in AGB and BGB allocations across treeline ecotones, which should be considered when estimating carbon dynamics with shifting treelines.