Jin-Sheng He

Community assembly during secondary forest succession in a Chinese subtropical forest

Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30 × 30 m plots (n = 27) from five successional stages ( 1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10 × 10 m quadrant of each plot. In addition, we measured a number of environmen...

Stoichiometry and large-scale patterns of leaf carbon and nitrogen in the grassland biomes of China

Nitrogen (N) and carbon–nitrogen (C:N) ratio are key foliar traits with great ecological importance, but their patterns across biomes have only recently been explored. We conducted a systematic census of foliar C, N and C:N ratio for 213 species, from 41 families over 199 research sites across the grassland biomes of China following the same protocol, to explore how different environmental conditions and species composition affect leaf N and C:N stoichiometry. Leaf C:N stoichiometry is stable in three distinct climatic regions in Inner Mongolia, the Tibetan Plateau, and Xinjiang Autonomous Region, despite considerable variations among co-existing species and among different vegetation types. Our results also show that life form and genus identity explain more than 70% of total variations of foliar N and C:N ratio, while mean growing season temperature and growing season precipitation explained only less than 3%. This suggests that, at the biome scale, temperature affects leaf N mainly through a change in plant species composition rather than via temperature itself. When our data were pooled with a global dataset, the previously observed positive correlation between leaf N and mean annual temperature (MAT) at very low MATs, disappeared. Thus, our data do not support the previously proposed biogeochemical hypothesis that low temperature limitations on mineralization of organic matter and N availability in soils lead to low leaf N in cold environments.

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

The links between ecosystem multifunctionality and above- and belowground biodiversity are mediated by climate

Plant biodiversity is often correlated with ecosystem functioning in terrestrial ecosystems. However, we know little about the relative and combined effects of above- and belowground biodiversity on multiple ecosystem functions (for example, ecosystem multifunctionality, EMF) or how climate might mediate those relationships. Here we tease apart the effects of biotic and abiotic factors, both above- and belowground, on EMF on the Tibetan Plateau, China. We found that a suite of biotic and abiotic variables account for up to 86% of the variation in EMF, with the combined effects of above- and belowground biodiversity accounting for 45% of the variation in EMF. Our results have two important implications: first, including belowground biodiversity in models can improve the ability to explain and predict EMF. Second, regional-scale variation in climate, and perhaps climate change, can determine, or at least modify, the effects of biodiversity on EMF in natural ecosystems.

Taxonomic, phylogenetic, and environmental trade-offs between leaf productivity and persistence

Assessing the influence of climate, soil fertility, and species identity on leaf trait relationships is crucial for understanding the adaptations of plants to their environment and for interpreting leaf trait relationships across spatial scales. In a comparative field study of 171 plant species in 174 grassland sites across China, we examined the trade-offs, defined as negative covariance between two traits, between leaf persistence (leaf mass per area, LMA) and leaf productivity (mass-based photosynthetic rate, Amass, N and P content, and photosynthetic N use efficiency, PNUE). We asked to which extent these trade-offs were influenced by: (1) variation among sites within species, decomposed into variation due to climatic and soil variables; (2) variation among species within sites, decomposed into variation among taxonomic, functional, or phylogenetic groups; and (3) the joint contribution of variation among species and sites. We used mixed-model analysis of covariance to partition bivariate relationships between leaf traits into trade-off components. We found significant mass-based persistence-productivity trade-offs of LMA-Amass, LMA-N, LMA-P, and LMA-PNUE consistent with previous broadscale findings. Overall, (1) variation among sites within species explained 14-23%, (2) variation among species within sites explained 20-34%, and (3) the two together explained 42-63% of the total covariance between leaf traits. Interspecific trade-offs of LMA-Amass, LMA-N, and LMA-P were stronger than inter-site ones. A relatively low amount of covariance was explained by climatic and soil variables. However, we found the trade-offs were stronger for LMA-N and LMA-P at higher precipitation and for LMA-PNUE at greater soil fertility, if displayed by major axis regression, which combined both intra- and interspecific variation. Residual trade-offs within species and sites were weak, suggesting that intraspecific, intra-site variation in physiology was less important than variation imposed by species identity or environmental differences among sites. Our results from grassland biomes add evidence for the fundamental nature of productivity-persistence trade-offs in plants. No individual factor emerged as the single major cause for these trade-offs. Rather, the total covariance between leaf traits was explained by a combination of factors, each contributing a range of explanatory power.

Soil organic carbon pools and stocks in permafrost-affected soils on the tibetan plateau.

The Tibetan Plateau reacts particularly sensitively to possible effects of climate change. Approximately two thirds of the total area is affected by permafrost. To get a better understanding of the role of permafrost on soil organic carbon pools and stocks, investigations were carried out including both discontinuous (site Huashixia, HUA) and continuous permafrost (site Wudaoliang, WUD). Three organic carbon fractions were isolated using density separation combined with ultrasonic dispersion: the light fractions (<1.6 g cm−3) of free particulate organic matter (FPOM) and occluded particulate organic matter (OPOM), plus a heavy fraction (>1.6 g cm−3) of mineral associated organic matter (MOM). The fractions were analyzed for C, N, and their portion of organic C. FPOM contained an average SOC content of 252 g kg−1. Higher SOC contents (320 g kg−1) were found in OPOM while MOM had the lowest SOC contents (29 g kg−1). Due to their lower density the easily decomposable fractions FPOM and OPOM contribute 27% (HUA) and 22% (WUD) to the total SOC stocks. In HUA mean SOC stocks (0–30 cm depth) account for 10.4 kg m−2, compared to 3.4 kg m−2 in WUD. 53% of the SOC is stored in the upper 10 cm in WUD, in HUA only 39%. Highest POM values of 36% occurred in profiles with high soil moisture content. SOC stocks, soil moisture and active layer thickness correlated strongly in discontinuous permafrost while no correlation between SOC stocks and active layer thickness and only a weak relation between soil moisture and SOC stocks could be found in continuous permafrost. Consequently, permafrost-affected soils in discontinuous permafrost environments are susceptible to soil moisture changes due to alterations in quantity and seasonal distribution of precipitation, increasing temperature and therefore evaporation.

Leaf‐Level Physiology, Biomass, and Reproduction of Phytolacca americana under Conditions of Elevated CO2 and Altered Temperature Regimes

The effects of increasing air temperature and changing daily temperature regime under conditions of elevated CO2 on the physiology, biomass, and reproduction of a C3 plant species were investigated. Phytolacca americana L. (Phytolaccaceae) was grown under either ambient (370 μmol mol−1) or elevated (700 μmol mol−1) CO2 at three air temperature regimes (day/night temperatures of 26°/20°C, T1; 30°/24°C, T2; and 28°/24°C, T3). Length of day/night temperature exposure was adjusted so that average daily temperature was 22°C in T1 and 26°C in T2 and T3. Daily temperature regime was different for T2 and T3: plants in T2 experienced a higher maximum daily temperature but for a shorter daily duration than plants in T3. Elevated CO2 increased photosynthetic rate, total biomass, and root‐to‐shoot ratio (RSR) and decreased stomatal conductance and transpiration as well as allocation to reproduction. In contrast, elevated temperatures had no effect on photosynthetic rate, stomatal conductance, or total biomass, but they decreased RSR and increased transpiration, reproductive biomass, and allocation. Both elevated CO2 and increased temperatures advanced timing of flowering. The plant‐level transpiration rate exhibited a unique response to each of the daily temperature regime treatments. These results indicate that elevated CO2 and increased temperatures elicit different responses at the physiological and whole‐plant levels in P. americana, with little interaction between the CO2 and temperature effects. Furthermore, some evidence indicates that a changing daily temperature regime may be an important factor determining plant responses to warming temperatures and should be incorporated into predictions of plant and ecosystem responses to future climate change.

Taxonomic identity, phylogeny, climate and soil fertility as drivers of leaf traits across Chinese grassland biomes

Although broad-scale inter-specific patterns of leaf traits are influenced by climate, soil, and taxonomic identity, integrated assessments of these drivers remain rare. Here, we quantify these drivers in a field study of 171 plant species in 174 sites across Chinese grasslands, including the Tibetan Plateau, Inner Mongolia, and Xinjiang. General linear models were used to partition leaf trait variation. Of the total variation in leaf traits, on average 27% is due to taxonomic or phylogenetic differences among species within sites (pure species effect), 29% to variation among sites within species (pure site effect), 38% to joint effects of taxonomic and environmental factors (shared effect), and 6.2% to within-site and within-species variation. Examining the pure site effect, climate explained 7.8%, soil explained 7.4%, and climate and soil variables together accounted for 11%, leaving 18% of the inter-site variation due to factors other than climate or soil. The results do not support the hypothesis that soil fertility is the “missing link” to explain leaf trait variation unexplained by climatic factors. Climate- and soil-induced leaf adaptations occur mostly among species, and leaf traits vary little within species in Chinese grassland plants, despite strongly varying climate and soil conditions.

On the combined effect of soil fertility and topography on tree growth in subtropical forest ecosystems—a study from SE China

Aims The aim of our research was to understand small-scale effects of topography and soil fertility on tree growth in a forest biodiversity and ecosystem functioning (BEF) experiment in subtropical SE China. Methods Geomorphometric terrain analyses were carried out at a spatial resolution of 5×5 m. Soil samples of different depth increments and data on tree height were collected from a total of 566 plots (667 m2 each). The soils were analyzed for carbon (soil organic carbon [SOC]), nitrogen, acidity, cation exchange capacity (CEC), exchangeable cations and base saturation as soil fertility attributes. All plots were classified into geomorphological units. Analyses of variance and linear regressions were applied to all terrain, soil fertility and tree growth attributes. Important Findings In general, young and shallow soils and relatively small differences in stable soil properties suggest that soil erosion has truncated the soils to a large extent over the whole area of the experiment. This explains the concurrently increasing CEC and SOC stocks downslope, in hollows and in valleys. However, colluvial, carbon-rich sediments are missing widely due to the convexity of the footslopes caused by uplift and removal of eroded sediments by adjacent waterways. The results showed that soil fertility is mainly influenced by topography. Monte–Carlo flow accumulation (MCCA), curvature, slope and aspect significantly affected soil fertility. Furthermore, soil fertility was affected by the different geomorphological positions on the experimental sites with ridge and spur positions showing lower exchangeable base cation contents, especially potassium (K), due to leaching. This geomorphological effect of soil fertility is most pronounced in the topsoil and decreases when considering the subsoil down to 50cm depth. Few soil fertility attributes affect tree height after 1–2 years of growth, among which C stocks proved to be most important while pHKCl and CEC only played minor roles. Nevertheless, soil acidity and a high proportion of Al on the exchange complex affected tree height even after only 1–2 years growth. Hence, our study showed that forest nutrition is coupled to a recycling of litter nutrients, and does not only depend on subsequent supply of nutrients from the mineral soil. Besides soil fertility, topography affected tree height. We found that especially MCCA as indicator of water availability affected tree growth at small-scale, as well as aspect. Overall, our synthesis on the interrelation between fertility, topography and tree growth in a subtropical forest ecosystem in SE China showed that topographic heterogeneity lead to ecological gradients across geomorphological positions. In this respect, small-scale soil–plant interactions in a young forest can serve as a driver for the future development of vegetation and biodiversity control on soil fertility. In addition, it shows that terrain attributes should be accounted for in ecological research.

Climate warming reduces the temporal stability of plant community biomass production

Anthropogenic climate change has emerged as a critical environmental problem, prompting frequent investigations into its consequences for various ecological systems. Few studies, however, have explored the effect of climate change on ecological stability and the underlying mechanisms. We conduct a field experiment to assess the influence of warming and altered precipitation on the temporal stability of plant community biomass in an alpine grassland located on the Tibetan Plateau. We find that whereas precipitation alteration does not influence biomass temporal stability, warming lowers stability through reducing the degree of species asynchrony. Importantly, biomass temporal stability is not influenced by plant species diversity, but is largely determined by the temporal stability of dominant species and asynchronous population dynamics among the coexisting species. Our findings suggest that ongoing and future climate change may alter stability properties of ecological communities, potentially hindering their ability to provide ecosystem services for humanity.