东北东部5种温带森林的春季土壤呼吸

春季土壤解冻过程是中高纬度地区森林生态系统土壤呼吸(即土壤表面CO2通量,RS)年内变化的一个关键时期,但此期间RS的时间动态规律及其控制机理尚不清楚。以我国东北东部5种温带森林为研究对象,采用静态箱-气相色谱法测定春季土壤解冻时期RS动态及其相关的环境因子。结果表明:在土壤解冻过程中,RS受林型、解冻时期及其交互作用的显著影响。红松(Pinus koraiensis)林、落叶松(Larix gmelinii)林、硬阔叶林、杨桦(Populus davidiana-Betulaplaty phylla)林和蒙古栎(Quercus mongolica)林的RS变化范围依次为:10.0196.0mg·m-·2h-1,5.8217.1mg·m-·2h-1,9.7382.1mg·m-·2h-1,15.8-269.0mg·m-·2h-1和35.9262.5mg·m-·2h-1。RS的平均值随着解冻的进程而增大,其变化趋势大致与土壤温度的变化相吻合。土壤温度极显著地影响RS(R2=0.46-0.77),而土壤含水量对RS的影响则因林型和土壤深度而异。5种林型的土壤呼吸温度系数(Q10)依次为:落叶松林10.9,硬阔叶林7.1,红松林6.5,杨桦林4.3和蒙古栎林2.3。进一步的研究应该集中研究春季自然解冻过程中土壤呼吸的控制机制,尤其是土壤呼吸与土壤微生物种群动态及其活性之间的关系。     

Seasonality and drivers of stem CO2 efflux for four temperate coniferous tree species

Aims Stem CO₂ efflux (E_s) is an important component of annual carbon budget in forest ecosystems, but how biotic and environmental factors regulate seasonal and inter-specific variations in E_s is poorly understood. The objectives of this study were: (1) to compare seasonal dynamics in E_s for four temperate coniferous tree species in northeastern China, including Korean pine (Pinus koraiensis), Korean spruce (Picea koraiensis), Mongolian pine (Pinus sylvestris var. mongolica), and Dahurian larch (Larix gmelinii); and (2) to explore factors driving the inter-specific variability in E_s during the growing and non-growing seasons.
Methods Ten to twelve trees for each tree species were sampled for E_s and stem temperature at 1 cm depth beneath the bark (T_s) measurements in situ with an infrared gas analyzer (LI-6400 IRGA) and a digital thermometer, respectively, from July to October 2013 and March to July 2014. The daily stem circumference increment (S_i), sapwood nitrogen concentration ([N]), and related environmental factors were monitored simultaneously.
Important findings The temporal variation in E_s for the four tree species overall followed the changes in T_s throughout the study period, with the maxima occurring in the summer months (late May to early July) characterized by higher temperature and more rapid stem growth and the minima in spring (late March to April) or autumn (October) having lower temperature. T_s accounted for 42%-91% and 56%-89% of variations in E_s during the growing (May to September) and non-growing (other months) seasons, respectively. Furthermore, apart from T_s, we also found significant regression relationships between E_s and S_i, relative air humidity and [N] during the growing season, but their forms and correlation coefficients were species-dependent. These results indicated that T_s was the dominant environmental factor affecting seasonal variations in E_s, but the magnitude of the effect varied with tree species and growth rhythm. Mean E_s for each of the four tree species was significantly higher in the growing season than in the non-growing season, whereas within the season there were also significant differences in mean E_s among the tree species (all p < 0.05). The temperature sensitivity of E_s (Q₁₀ value) did not differ significantly among the tree species during the growing season, ranging from 1.64 for Dahurian larch to 2.09 for Mongolian pine, but did differ during the non-growing season which varied from 1.80 for Korean pine to 3.14 for Dahurian larch. Moreover, Korean spruce, Mongolian pine and Dahurian larch had significantly greater Q₁₀ values in the non-growing season than in the growing season (p < 0.05). These findings suggested that the differences of the response of E_s to temperature change for different tree species were mainly from the non-growing season. Because the seasonality and inter-specific variability in E_s for these temperate coniferous tree species were primarily controlled by multiple factors such as temperature, we conclude that using a single annual temperature response curve to estimate the annual E_s may lead to more uncertainty.     

Decomposing Spatial β‐Diversity in the temperate forests of Northeastern China

β‐Diversity, which describes the extent of change in species composition in a given region, has become a core issue in ecology in recent years. However, it is hard to understand the underlying mechanisms of β‐diversity by using indices that yield identical values under species replacement and nestedness pattern. Partitioning β‐diversity into turnover (caused by species replacement among plots) and nestedness components (caused by species loss or gain among plots) may provide improved understanding of the variation in species composition. Here, we collected presence–absence data of 456 one‐tenth ha circular plots in the temperate forests of Northeastern China spanning a latitudinal range of 12° (41–53°N). We decomposed β‐diversity to assess the relative contribution of the turnover and nestedness components across latitudinal gradients. We used regression analysis to assess the relationship between spatial distance and β‐diversity. We applied variation partitioning to evaluate the importance of the measured environmental and spatial variables in explaining β‐diversity. We used the Tukey honest significant difference test to test the differences of β‐diversity along latitudinal gradients. Pearson correlations (r) and significance (p‐value) were computed using the Mantel tests to verify the relationship between distance and β‐diversity. The ANOVA test was used to verify whether the variation of β‐diversity explained by the environment and distance was significant. Our results showed that (1) β‐diversity and the turnover component were higher at low latitudes (zones A and B) than at high latitudes (zones C and D), while there was no relationship between the nestedness component and latitude. (2) The turnover component was dominant. (3) The spatial distance explained more variation of β‐diversity than the measured environmental factors. Therefore, we conclude that β‐diversity is mainly a product of species turnover in our temperate forests, suggesting that different localities harbor different species. We find that decomposing β‐diversity into the turnover and nestedness components is a useful approach to explore the variation of community composition and their causes.     

Temporal and spatial variations of stem CO2 efflux of three species in subtropical China

Aims Although stem CO₂ efflux is critical to ecosystem carbon and energy balance and its feedback to future climate change, little information is available on stem CO₂ efflux and its responses to temperature, especially in subtropical China. This study aims to (i) evaluate the temporal and spatial variations of stem CO₂ efflux of three species, including oak (Quercus acutissima Carr.), masson pine (Pinus massoniana Lamb.) and loblolly pine (Pinus taeda Linn.) in subtropical China and (ii) analyze the temperature sensitivity of stem CO₂ efflux in the three species based on 2-year field measurements.Methods We measured stem CO₂ efflux and stem temperature (at 3 cm depth) of the three species using the horizontally oriented soil chamber technique from September 2008 to August 2010. We also conducted a 24-h measurement to examine the diurnal variation of stem CO₂ efflux in three consecutive days in April 2009.Important findings The temporal dynamics of stem CO₂ efflux followed the change of the stem temperature in a 3-cm depth with a bell-shaped curve in the three species. Stem temperature explained 77–85% of the seasonal variations of stem CO₂ efflux over the entire study period in the three species. The temperature sensitivity (Q 10) of stem CO₂ efflux was obviously different among the three species with higher Q 10 value found in oak (2.24) and lower values in the coniferous species (1.76 and 1.63). Our results also showed that the Q 10 values of stem CO₂ efflux in all the three species were lower in the growing season than that in the non-growing season, indicating that the growth and maintenance respiration had different temperature responses. Moreover, we found that the temperature-normalized stem CO₂ efflux (R 10) changed greatly between the growing and non-growing seasons in oak and masson pine, but not in loblolly pine. Additionally, we also found that in the non-growing season, the principal factor responsible for the spatial variation of stem CO₂ efflux among the 15 sampling trees was sapwood volume, whereas in the growing season, stem CO₂ efflux was closely related to annual dry-matter production in the three subtropical species.     

产于中国西南高山温带森林的二个棒瑚菌新种

本文根据形态学特征和分子系统学分析报道了产于中国西南山地温带森林的棒瑚菌属新种2个。第一个新种金黄棒瑚菌Clavariadelphus aurantiacus具有亮橙黄色的子实体和不育的子实体顶部,第二个新种纤细棒瑚菌C. tenuis具有细长的子实体并且菌丝不具锁状联合。为两个新种提供了详细的描述、绘制了黑白线条图、拍摄了彩色照片,并且与类似的物种进行了比较。运用核糖体转录区间序列分析了棒瑚菌属的系统发育关系。提供了中国已知棒瑚菌的检索表。     

Stem CO2 efflux in six co‐occurring tree species: underlying factors and ecological implications

Stem respiration plays a role in species coexistence and forest dynamics. Here we examined the intra‐ and inter‐specific variability of stem CO₂ efflux (E) in dominant and suppressed trees of six deciduous species in a mixed forest stand: Fagus sylvatica L., Quercus petraea [Matt.] Liebl, Quercus pyrenaica Willd., Prunus avium L., Sorbus aucuparia L. and Crataegus monogyna Jacq. We conducted measurements in late autumn. Within species, dominants had higher E per unit stem surface area (E_s) mainly because sapwood depth was higher than in suppressed trees. Across species, however, differences in E_s corresponded with differences in the proportion of living parenchyma in sapwood and concentration of non‐structural carbohydrates (NSC). Across species, E_s was strongly and NSC marginally positively related with an index of drought tolerance, suggesting that slow growth of drought‐tolerant trees is related to higher NSC concentration and E_s. We conclude that, during the leafless period, E is indicative of maintenance respiration and is related with some ecological characteristics of the species, such as drought resistance; that sapwood depth is the main factor explaining variability in E_s within species; and that the proportion of NSC in the sapwood is the main factor behind variability in E_s among species.     

Soil CO2 efflux in response to the addition of water and fertilizer in temperate semiarid grassland in northern China

Background and aims

Knowledge about the effects of water and fertilizer on soil CO₂ efflux (SCE) and Q ₁₀ is essential for understanding carbon (C) cycles and for evaluating future global C balance. A two-year field experiment was conducted to determine the effects of water, fertilizer, and temperature on SCE in semiarid grassland in northern China.

Methods

SCE, as well as environmental factors was measured in two grasslands, one with bunge needlegrass (BNE, Stipa bungeana) and one with purple alfalfa (ALF, Medicago sativa), with four treatments: CK (unwatered and unfertilized); W (50 mm water addition yr^?1); F (50 kg phosphorus (P) fertilizer ha^?1 yr^?1 for ALF, 100 kg nitrogen (N)?+?50 kg P fertilizer ha^?1 yr^?1 for BNE); and W + F.

Results

During the 11-month experimental period from July 2010 to October 2011, the addition of water consistently stimulated mean SCE in BNE and ALF, and the positive effects were relatively stronger during dry seasons. P fertilization consistently enhanced SCE in ALF, and the positive effect was strongly dependent on the availability of soil water. The effects of N plus P fertilization on SCE in BNE varied seasonally from significant increases to small reductions to no response. Water addition increased the Q ₁₀ of SCE in ALF by 11 % but had no effect in BNE. Fertilization, however, reduced the Q ₁₀ of SCE by 21 % and 13 % for BNE and ALF, respectively. Models that rely only on Q ₁₀ underestimated the emissions of soil CO₂ by 8–15 % at the study site, which was mediated by species and treatment.

Conclusions

Responses of SCE and its temperature sensitivity to water and fertilizer may vary with species and depend on the period of measurement. Models of SCE need to incorporate the availability of ecosystemic water and nutrients, as well as species, and incorporate different environmental factors when determining the impact of water, nutrients, and species on SCE.     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

Temporal patterns of soil CO2 efflux in a temperate Korean Larch (Larix olgensis Herry.) plantation, Northeast China

There is little information available regarding seasonal and annual variations in soil CO₂ efflux from Korean Larch plantations, which are an important component of forests’ carbon balance in temperate China. In this study, the soil respiration rate (R _s), soil temperature (T ₁₀) and soil moisture (SM₁₀) at 10 cm depth were observed in a Korean Larch (Larix olgensis Herry.) plantation in Northeast China from 2008 to 2012. Mean R _s in growing season (GS) varied greatly, ranged from 2.32 ± 0.08 to 3.88 ± 0.09 μmol CO₂ m^?2 s^?1 (mean ± SE) over the period of 2008–2012. In comparison with T-model, the increase of explained variability by applying both T ₁₀ and SM₁₀ to the T-M model is very small. It is indicated that R _s was controlled largely by T ₁₀ in the present study. By accounting for 22.2 and 17.7 % of the total soil CO₂ emissions in 2010/2011 and 2011/2012, respectively, the soil CO₂ efflux in dormant season (DS) was an essential component of the total soil CO₂ efflux. The Q ₁₀ value in the study period was always smaller for GS than DS, suggesting that soil carbon cycling may be more sensitive to the temperature changes at low than at high temperature range. These results indicated that climate changes may have great potential impacts on temperate Larch plantations in Northeast China, owing to soil carbon emissions of Larch plantation during the long period of DS being more sensitive to T ₁₀ than in GS, and played a significant role in the annual forest ecosystems carbon budget.     

东北东部森林生态系统土壤呼吸组分的分离量化

对森林生态系统的土壤呼吸组分进行分离和量化,确定不同组分CO2释放速率的控制因子,是估测局域和区域森林生态系统碳平衡研究中必不可少的内容。采用挖壕法和红外气体分析法测定无根和有根样地的土壤表面CO2通量(RS),确定东北东部6种典型森林生态系统RS中异养呼吸(RH)和根系自养呼吸(RA)的贡献量及其影响因子。具体研究目标包括:(1)量化各种生态系统的RH及其与主要环境影响因子的关系;(2)量化各种生态系统RS中根系呼吸贡献率(RC)的季节动态;(3)比较6种森林生态系统RH和RA的年通量。土壤温度、土壤含水量及其交互作用显著地影响森林生态系统的RH(R2=0.465~0.788),但其影响程度因森林生态系统类型而异。硬阔叶林和落叶松人工林的RH主要受土壤温度控制,其他生态系统RH受土壤温度和含水量的联合影响。各个森林生态系统类型的RC变化范围依次为:硬阔叶林32.40%~51.44%;杨桦林39.72%~46.65%;杂木林17.94%~47.74%;蒙古栎林34.31%~37.36%;红松人工林33.78%~37.02%;落叶松人工林14.39%~35.75%。每个生态系统类型RH年通量都显著高于RA年通量,其变化范围分别为337~540 gC.m-2.a-1和88~331 gC.m-2.a-1。不同生态系统间的RH和RA也存在着显著性差异。     

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

Rapid turnover of DOC in temperate forests accounts for increased CO2 production at elevated temperatures

The evidence for the contribution of soil warming to changes in atmospheric CO₂ concentrations and carbon stocks of temperate forest ecosystems is equivocal. Here, we use data from a beech/oak forest on concentrations and stable isotope ratios of dissolved organic carbon (DOC), phosphate buffer-extractable organic carbon, soil organic carbon (SOC), respiration and microbial gross assimilation of N to show that respired soil carbon originated from DOC. However, the respiration was not dependent on the DOC concentration but exceeded the daily DOC pool three to four times, suggesting that DOC was turned over several times per day. A mass flow model helped to calculate that a maximum of 40% of the daily DOC production was derived from SOC and to demonstrate that degradation of SOC is limiting respiration of DOC. The carbon flow model on SOC, DOC, microbial C mobilization/immobilization and respiration is linked by temperature-dependent microbial and enzyme activity to global warming effects of CO₂ emitted to the atmosphere.     

Spatial pattern of diversity in an old-growth temperate forest in Northeastern China

Species diversity has attracted particular attention because of its significance for helping determine present species performance and likely future community composition. The spatial pattern of species diversity (species richness, abundance and Shannon diversity) in Changbai temperate forest in Northeastern China was studied to investigate the present and likely causes for the formation of spatial patterns. To fulfill this goal, three aspects of diversity were addressed: 1) changes in the relationships of the diversity variables, species richness, abundance and Shannon diversity, to sampling area and sampling design. The three diversity variables were found to respond to sampling area in a dissimilar way. Sampling design had no significant effect on the diversity variable-area curves. The power function, which was derived under the assumption that the forest was in equilibrium, did not fit the observed species-area curves, indicating that the Changbai temperate forest was probably not in equilibrium. 2) Variograms, used to examine the spatial structure of species diversity, showed that the spatial structure of species diversity in the Changbai temperate forest was weakly anisotropic. 3) Partitioning the variation of species diversity into spatial and environmental factors indicated that the spatial pattern of the Changbai forest community was unpredictable, probably because there were many undetermined processes controlling its development.     

Germinative behaviour of ten tree species in white-water floodplain forests in central Amazonia

Amazonian floodplain forests (known as várzea) are classified into high or low várzea depending on the spatial position on the plains. This topographic feature exposes the terrain over different time periods of inundation, causing a major limiting factor for tree seedling establishment. We hypothesize that, strategically, most of the seeds produced by trees in low várzea forests germinate faster and in synchrony (temporally concentrated germination), and that their seedlings tend to have cotyledons without reserve or foliaceous cotyledons (PEF). By contrast, seeds produced by high-várzea specialist trees exhibit slower and temporally scattered germination, and their seedlings tend to have reserve storage cotyledons (CHR). Generalist species may show no clear pattern or may be related to high-várzea species. To test this hypothesis, diaspores of 10 tree species were collected: five of low-várzea specialist trees, three of high-várzea specialist trees and two of generalist species. Seedling emergence and morphology were monitored daily in a nursery for a period of 150 days of being subjected to non-flooded (sown directly in várzea soil) and flooded conditions (15 days in water before sowing in the same soil). The seedling emergence of low-várzea species showed an increase of 37% in germinability whereas high-várzea and generalist species exhibited a decrease of 38% and 35% of germinability, respectively. Foliaceous cotyledons were preferentially found in seedlings of low-várzea species, and storage cotyledons were more common in those of high-várzea species, indicating how cotyledon morphology may determine the amount and use of resources available to a seedling during the first stages of establishment and growth. Conservation plans aiming for the maintenance of ecosystem services must consider these strategies.     

Effects of warming on seed germination of woody species in temperate secondary forests

• Seed germination, a critical stage of the plant life cycle providing a link between seeds and seedlings, is commonly temperature-dependent. The global average surface temperature is expected to rise, but little is known about the responses of seed germination of woody plants in temperate forests to warming.
• In the present study, dried seeds of 23 common woody species in temperate secondary forests were incubated at three temperature sequences without cold stratification and after experiencing cold stratification. We calculated five seed germination indices and the comprehensive membership function value that summarized the above indicators.
• Compared to the control, +2 and +4 °C treatments without cold stratification shortened germination time by 14% and 16% and increased the germination index by 17% and 26%, respectively. For stratified seeds, +4 °C treatment increased germination percentage by 49%, and +4 and +2 °C treatments increased duration of germination and the germination index, and shortened mean germination time by 69%, 458%, 29% and 68%, 110%, 12%, respectively. The germination of Fraxinus rhynchophylla and Larix kaempferi were most sensitive to warming without and with cold stratification, respectively. Seed germination of shrubs was the least sensitive to warming among functional types.
• These findings indicate warming (especially extreme warming) will enhance the seedling recruitment of temperate woody species, primarily via shortening the germination time, particularly for seeds that have undergone cold stratification. In addition, shrubs might narrow their distribution range

     

Soil CO2 efflux in uneven-aged managed forests: temporal patterns following harvest and effects of edaphic heterogeneity

Increased nutrient availability can have a large impact in Australian woodland ecosystems, many of which are very poor in nutrients, particularly phosphorus. A study was conducted in an urban Banksia woodland remnant in Perth, southwest Western Australia to test the hypothesis that the soil nutrient status in areas in good condition (GC), poor condition invaded by the perennial grass Ehrharta calycina (PCe), and poor condition invaded by the perennial herb Pelargonium capitatum (PCp), is reflected in the nutrient status of the native and introduced species. Leaf concentrations of P, K, N, Na, Ca, Mg, S, B, Cu, Fe, Mn and Zn of six native (Banksia attenuata, Banksia menziesii, Allocasuarina humilis, Melaleuca systena, Macrozamia fraseri and Conostylis aculeata) and four introduced species (Ehrharta calycina, Pelargonium capitatum, Gladiolus caryophyllaceus and Briza maxima), were measured. Soil pH, electrical conductivity, N (total), P (total), available P, K, S and organic C were assessed beneath all species on all sites. Significantly higher concentrations of soil P (total) and P (HCO₃) were found at PCe and PCp sites than GC sites, while PCp sites also had significantly higher soil concentrations for N (total) and S. Principal Components Analysis of the leaf analyses showed (a) individual species have characteristically different nutrient concentrations; (b) the introduced species Ehrharta calycina and Pelargonium capitatum clustered separately from each other and by vegetation condition. Leaf concentrations of P were significantly (P<0.05) higher, and K and Cu were significantly lower in PCe and PCp sites compared with those at GC sites. Introduced species leaf nutrient concentrations were significantly greater than native species for all nutrients except Mn which was significantly lower, with no differences for Mg and B. The results indicate a key role for P in the Banksia woodland, and we conclude that higher levels of available P at invaded sites are having a detrimental impact on the ecosystem. These results provide new knowledge to enhance conservation practices for the management of the key threatening process of invasion within a biodiversity hot spot. Section Editor: T. Kalapos     

Determinants of invasion by single versus multiple plant species in temperate lowland forests

Biological Invasions - Invasions of alien plants pose a serious threat to native biodiversity and ecosystem processes. Forests are considered more resistant to invasion due to limited light...