A Synthesis of Climate and Vegetation Cover Effects on Biogeochemical Cycling in Shrub-Dominated Drylands

Semi-arid and arid ecosystems dominated by shrubs (“dry shrublands”) are an important component of the global C cycle, but impacts of climate change and elevated atmospheric CO₂ on biogeochemical cycling in these ecosystems have not been synthetically assessed. This study synthesizes data from manipulative studies and from studies contrasting ecosystem processes in different vegetation microsites (that is, shrub or herbaceous canopy versus intercanopy microsites), to assess how changes in climate and atmospheric CO₂ affect biogeochemical cycles by altering plant and microbial physiology and ecosystem structure. Further, we explore how ecosystem structure impacts on biogeochemical cycles differ across a climate gradient. We found that: (1) our ability to project ecological responses to changes in climate and atmospheric CO₂ is limited by a dearth of manipulative studies, and by a lack of measurements in those studies that can explain biogeochemical changes, (2) changes in ecosystem structure will impact biogeochemical cycling, with decreasing pools and fluxes of C and N if vegetation canopy microsites were to decline, and (3) differences in biogeochemical cycling between microsites are predictable with a simple aridity index (MAP/MAT), where the relative difference in pools and fluxes of C and N between vegetation canopy and intercanopy microsites is positively correlated with aridity. We conclude that if climate change alters ecosystem structure, it will strongly impact biogeochemical cycles, with increasing aridity leading to greater heterogeneity in biogeochemical cycling among microsites. Additional long-term manipulative experiments situated across dry shrublands are required to better predict climate change impacts on biogeochemical cycling in deserts.     

Determinants of soil organic carbon sequestration and its contribution to ecosystem carbon sinks of planted forests

The area of forest established through afforestation/reforestation has been increasing on a global scale, which is particularly important as these planted forests attenuate climate change by sequestering carbon. However, the determinants of soil organic carbon (SOC) sequestration and their contribution to the ecosystem carbon sink of planted forests remain uncertain. By using globally distributed data extracted from 154 peer‐reviewed publications and a total of 355 sampling points, we investigated above‐ground biomass carbon (ABC) sequestration and SOC sequestration across three different climatic zones (tropical, warm temperate, and cold temperate) through correlation analysis, regression models, and structural equation modeling (SEM). We found that the proportion of SOC sequestration in the ecosystem C sequestration averaged 14.1% globally, being the highest (27.0%) in the warm temperate and the lowest (10.7%) in the tropical climatic zones. The proportion was mainly affected by latitude. The sink rate of ABC (R_ABC) in tropical climates (2.48 Mg C ha^?1 year^?1) and the sink rate of SOC (R_SOC) in warm temperate climates (0.96 Mg C ha^?1 year^?1) were higher than other climatic zones. The main determinants of R_SOC were the number of frost‐free days, latitude, mean annual precipitation (MAP), and SOC density (SOCD) at the initial observation; however, these variables depended on the climatic zone. According to the SEM, frost‐free period, mean annual temperature (MAT) and MAP are the dominant driving factors affecting R_SOC in Chinese plantations. MAT has a positive effect on R_SOC, and global warming may increase R_SOC of temperate plantations in China. Our findings highlight the determinants of SOC sequestration and quantitatively reveal the substantial global contribution of SOC sequestration to ecosystem carbon sink provided by planted forests. Our results help managers identify and control key factors to increase carbon sequestration in forest ecosystems.     

Responses of dryland soil respiration and soil carbon pool size to abrupt vs. gradual and individual vs. combined changes in soil temperature, precipitation, and atmospheric [CO2]: a simulation analysis

With the large extent and great amount of soil carbon (C) storage, drylands play an important role in terrestrial C balance and feedbacks to climate change. Yet, how dryland soils respond to gradual and concomitant changes in multiple global change drivers [e.g., temperature (T_s), precipitation (Ppt), and atmospheric [CO₂] (CO₂)] has rarely been studied. We used a process‐based ecosystem model patch arid land simulator to simulate dryland soil respiration (R_s) and C pool size (C_s) changes to abrupt vs. gradual and single vs. combined alterations in T_s, Ppt and CO₂ at multiple treatment levels. Results showed that abrupt perturbations generally resulted in larger R_s and had longer differentiated impacts than did gradual perturbations. R_s was stimulated by increases in T_s, Ppt, and CO₂ in a nonlinear fashion (e.g., parabolically or asymptotically) but suppressed by Ppt reduction. Warming mainly stimulated heterotrophic R_s (i.e., R_h) whereas Ppt and CO₂ influenced autotrophic R_s (i.e., R_a). The combined effects of warming, Ppt, and CO₂ were nonadditive of primary single‐factor effects as a result of substantial interactions among these factors. Warming amplified the effects of both Ppt addition and CO₂ elevation whereas Ppt addition and CO₂ elevation counteracted with each other. Precipitation reduction either magnified or suppressed warming and CO₂ effects, depending on the magnitude of factor's alteration and the components of R_s (R_a or R_h) being examined. Overall, Ppt had dominant influence on dryland R_s and C_s over T_s and CO₂. Increasing Ppt individually or in combination with T_s and CO₂ benefited soil C sequestration. We therefore suggested that global change experimental studies for dryland ecosystems should focus more on the effects of precipitation regime changes and the combined effects of Ppt with other global change factors (e.g., T_s, CO₂, and N deposition).     

Net ecosystem CO2 exchange in Mojave Desert shrublands during the eighth year of exposure to elevated CO2

Arid ecosystems, which occupy about 35% of the Earth's terrestrial surface area, are believed to be among the most responsive to elevated [CO₂]. Net ecosystem CO₂ exchange (NEE) was measured in the eighth year of CO₂ enrichment at the Nevada Desert Free‐Air CO₂ Enrichment (FACE) Facility between the months of December 2003–December 2004. On most dates mean daily NEE (24 h) (μmol CO₂ m^?2 s^?1) of ecosystems exposed to elevated atmospheric CO₂ were similar to those maintained at current ambient CO₂ levels. However, on sampling dates following rains, mean daily NEEs of ecosystems exposed to elevated [CO₂] averaged 23 to 56% lower than mean daily NEEs of ecosystems maintained at ambient [CO₂]. Mean daily NEE varied seasonally across both CO₂ treatments, increasing from about 0.1 μmol CO₂ m^?2 s^?1 in December to a maximum of 0.5–0.6 μmol CO₂ m^?2 s^?1 in early spring. Maximum NEE in ecosystems exposed to elevated CO₂ occurred 1 month earlier than it did in ecosystems exposed to ambient CO₂, with declines in both treatments to lowest seasonal levels by early October (0.09±0.03 μmol CO₂ m^?2 s^?1), but then increasing to near peak levels in late October (0.36±0.08 μmol CO₂ m^?2 s^?1), November (0.28±0.03 μmol CO₂ m^?2 s^?1), and December (0.54±0.06 μmol CO₂ m^?2 s^?1). Seasonal patterns of mean daily NEE primarily resulted from larger seasonal fluctuations in rates of daytime net ecosystem CO₂ uptake which were closely tied to plant community phenology and precipitation. Photosynthesis in the autotrophic crust community (lichens, mosses, and free‐living cyanobacteria) following rains were probably responsible for the high NEEs observed in January, February, and late October 2004 when vascular plant photosynthesis was low. Both CO₂ treatments were net CO₂ sinks in 2004, but exposure to elevated CO₂ reduced CO₂ sink strength by 30% (positive net ecosystem productivity=127±17 g C m^?2 yr^?1 ambient CO₂ and 90±11 g C m^?2 yr^?1 elevated CO₂, P=0.011). This level of net C uptake rivals or exceeds levels observed in some forested and grassland ecosystems. Thus, the decrease in C sequestration seen in our study under elevated CO₂– along with the extensive coverage of arid and semi‐arid ecosystems globally – points to a significant drop in global C sequestration potential in the next several decades because of responses of heretofore overlooked dryland ecosystems.     

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.     

Canopy scale measurements of CO2 and water vapor exchange along a precipitation gradient in southern Africa

Short‐term measurements of carbon dioxide, water, and energy fluxes were collected at four locations along a mean annual precipitation gradient in southern Africa during the wet (growing) season with the purpose of determining how the observed vegetation–atmosphere exchange properties are functionally related to the long‐term climatic conditions. This research was conducted along the Kalahari Transect (KT), one in the global set of International Geosphere‐Biosphere Program transects, which covers a north–south aridity gradient, all on a homogenous sand formation. Eddy covariance instruments were deployed on a permanent tower in Mongu, Zambia (879 mm of rainfall per year), as well as on a portable tower in Maun (460 mm yr^?1), Okwa River Crossing (407 mm yr^?1), and Tshane (365 mm yr^?1), Botswana for several days at each site. The relationships between CO₂ flux, F_c, and photosynthetically active radiation were described well by a hyperbolic fit to the data at all locations except for Mongu, the wettest site. Here, there appeared to be an air temperature effect on F_c. While daytime values of F_c routinely approached or exceeded ?20 μmol m^?2 s^?1 at Mongu, the magnitude of F_c remained less than ?10 μmol m^?2 s^?1 when the air temperature was above 27°C. Canopy resistances to water vapor transfer, r_c, displayed an overall decline from the wetter sites to the more arid sites, but the differences in r_c could be almost exclusively accounted for by the decrease in leaf area index (LAI) from north to south along the KT. Ecosystem water use efficiency (WUE), defined as the ratio of net carbon flux to evapotranspiration, showed a general decrease with increasing vapor pressure deficit, D, for all of the sites. The magnitudes of WUE at a given D, however, were dissimilar for the individual sites and were found to be stratified according to the position of the sites along the long‐term aridity gradient. For example, Mongu, which has the wettest climate, has a much lower WUE for like levels of D than Tshane, which historically has the most arid climate. Given the similar inferred stomatal resistances between the sites, the disparate carbon uptake behavior for the grass vs. woody vegetation is the likely cause for the observed differences in WUE along the aridity gradient. The short‐term flux measurements provide a framework for evaluating the vegetation's functional adaptation to the long‐term climate and provide information that may be useful for predicting the dynamic response of the vegetation to future climate change.     

Variation in soil carbon stocks and their determinants across a precipitation gradient in West Africa

We examine the influence of climate, soil properties and vegetation characteristics on soil organic carbon (SOC) along a transect of West African ecosystems sampled across a precipitation gradient on contrasting soil types stretching from Ghana (15°N) to Mali (7°N). Our findings derive from a total of 1108 soil cores sampled over 14 permanent plots. The observed pattern in SOC stocks reflects the very different climatic conditions and contrasting soil properties existing along the latitudinal transect. The combined effects of these factors strongly influence vegetation structure. SOC stocks in the first 2 m of soil ranged from 20 Mg C ha^?1 for a Sahelian savanna in Mali to over 120 Mg C ha^?1 for a transitional forest in Ghana. The degree of interdependence between soil bulk density (SBD) and soil properties is highlighted by the strong negative relationships observed between SBD and SOC (r² > 0.84). A simple predictive function capable of encompassing the effect of climate, soil properties and vegetation type on SOC stocks showed that available water and sand content taken together could explain 0.84 and 0.86 of the total variability in SOC stocks observed to 0.3 and 1.0 m depth respectively. Used in combination with a suitable climatic parameter, sand content is a good predictor of SOC stored in highly weathered dry tropical ecosystems with arguably less confounding effects than provided by clay content. There was an increased contribution of resistant SOC to the total SOC pool for lower rainfall soils, this likely being the result of more frequent fire events in the grassier savannas of the more arid regions. This work provides new insights into the mechanisms determining the distribution of carbon storage in tropical soils and should contribute significantly to the development of robust predictive models of biogeochemical cycling and vegetation dynamics in tropical regions.     

Soil respiration is stimulated by elevated CO2 and reduced by summer drought: three years of measurements in a multifactor ecosystem manipulation experiment in a temperate heathland (CLIMAITE)

This study investigated the impact of predicted future climatic and atmospheric conditions on soil respiration (R_S) in a Danish Calluna‐Deschampsia‐heathland. A fully factorial in situ experiment with treatments of elevated atmospheric CO₂ (+130 ppm), raised soil temperature (+0.4 °C) and extended summer drought (5–8% precipitation exclusion) was established in 2005. The average R_S, observed in the control over 3 years of measurements (1.7 μmol CO₂ m^?2 sec^?1), increased 38% under elevated CO₂, irrespective of combination with the drought or temperature treatments. In contrast, extended summer drought decreased R_S by 14%, while elevated soil temperature did not affect R_S overall. A significant interaction between elevated temperature and drought resulted in further reduction of R_S when these treatments were combined. A detailed analysis of short‐term R_S dynamics associated with drought periods showed that R_S was reduced by ~50% and was strongly correlated with soil moisture during these events. Recovery of R_S to pre‐drought levels occurred within 2 weeks of rewetting; however, unexpected drought effects were observed several months after summer drought treatment in 2 of the 3 years, possibly due to reduced plant growth or changes in soil water holding capacity. An empirical model that predicts R_S from soil temperature, soil moisture and plant biomass was developed and accounted for 55% of the observed variability in R_S. The model predicted annual sums of R_S in 2006 and 2007, in the control, were 672 and 719 g C m^?2 y^?1, respectively. For the full treatment combination, i.e. the future climate scenario, the model predicted that soil respiratory C losses would increase by ~21% (140–150 g C m^?2 y^?1). Therefore, in the future climate, stimulation of C storage in plant biomass and litter must be in excess of 21% for this ecosystem to not suffer a reduction in net ecosystem exchange.     

Soil organic carbon dynamics jointly controlled by climate,carbon inputs,soil properties and soil carbon fractions

Soil organic carbon (SOC) dynamics are regulated by the complex interplay of climatic, edaphic and biotic conditions. However, the interrelation of SOC and these drivers and their potential connection networks are rarely assessed quantitatively. Using observations of SOC dynamics with detailed soil properties from 90 field trials at 28 sites under different agroecosystems across the Australian cropping regions, we investigated the direct and indirect effects of climate, soil properties, carbon (C) inputs and soil C pools (a total of 17 variables) on SOC change rate (r_C, Mg C ha^?1 yr^?1). Among these variables, we found that the most influential variables on r_C were the average C input amount and annual precipitation, and the total SOC stock at the beginning of the trials. Overall, C inputs (including C input amount and pasture frequency in the crop rotation system) accounted for 27% of the relative influence on r_C, followed by climate 25% (including precipitation and temperature), soil C pools 24% (including pool size and composition) and soil properties (such as cation exchange capacity, clay content, bulk density) 24%. Path analysis identified a network of intercorrelations of climate, soil properties, C inputs and soil C pools in determining r_C. The direct correlation of r_C with climate was significantly weakened if removing the effects of soil properties and C pools, and vice versa. These results reveal the relative importance of climate, soil properties, C inputs and C pools and their complex interconnections in regulating SOC dynamics. Ignorance of the impact of changes in soil properties, C pool composition and C input (quantity and quality) on SOC dynamics is likely one of the main sources of uncertainty in SOC predictions from the process‐based SOC models.     

Pathways regulating decreased soil respiration with warming in a biocrust‐dominated dryland

A positive soil carbon (C)‐climate feedback is embedded into the climatic models of the IPCC. However, recent global syntheses indicate that the temperature sensitivity of soil respiration (R_S) in drylands, the largest biome on Earth, is actually lower in warmed than in control plots. Consequently, soil C losses with future warming are expected to be low compared with other biomes. Nevertheless, the empirical basis for these global extrapolations is still poor in drylands, due to the low number of field experiments testing the pathways behind the long‐term responses of soil respiration (R_S) to warming. Importantly, global drylands are covered with biocrusts (communities formed by bryophytes, lichens, cyanobacteria, fungi, and bacteria), and thus, R_S responses to warming may be driven by both autotrophic and heterotrophic pathways. Here, we evaluated the effects of 8‐year experimental warming on R_S, and the different pathways involved, in a biocrust‐dominated dryland in southern Spain. We also assessed the overall impacts on soil organic C (SOC) accumulation over time. Across the years and biocrust cover levels, warming reduced R_S by 0.30 μmol CO₂ m^?2 s^?1 (95% CI = ?0.24 to 0.84), although the negative warming effects were only significant after 3 years of elevated temperatures in areas with low initial biocrust cover. We found support for different pathways regulating the warming‐induced reduction in R_S at areas with low (microbial thermal acclimation via reduced soil mass‐specific respiration and β‐glucosidase enzymatic activity) vs. high (microbial thermal acclimation jointly with a reduction in autotrophic respiration from decreased lichen cover) initial biocrust cover. Our 8‐year experimental study shows a reduction in soil respiration with warming and highlights that biocrusts should be explicitly included in modeling efforts aimed to quantify the soil C–climate feedback in drylands.     

A cross‐biome synthesis of soil respiration and its determinants under simulated precipitation changes

Soil respiration (R_s) is the second‐largest terrestrial carbon (C) flux. Although R_s has been extensively studied across a broad range of biomes, there is surprisingly little consensus on how the spatiotemporal patterns of R_s will be altered in a warming climate with changing precipitation regimes. Here, we present a global synthesis R_s data from studies that have manipulated precipitation in the field by collating studies from 113 increased precipitation treatments, 91 decreased precipitation treatments, and 14 prolonged drought treatments. Our meta‐analysis indicated that when the increased precipitation treatments were normalized to 28% above the ambient level, the soil moisture, R_s, and the temperature sensitivity (Q₁₀) values increased by an average of 17%, 16%, and 6%, respectively, and the soil temperature decreased by ?1.3%. The greatest increases in R_s and Q₁₀ were observed in arid areas, and the stimulation rates decreased with increases in climate humidity. When the decreased precipitation treatments were normalized to 28% below the ambient level, the soil moisture and R_s values decreased by an average of ?14% and ?17%, respectively, and the soil temperature and Q₁₀ values were not altered. The reductions in soil moisture tended to be greater in more humid areas. Prolonged drought without alterations in the amount of precipitation reduced the soil moisture and R_s by ?12% and ?6%, respectively, but did not alter Q₁₀. Overall, our synthesis suggests that soil moisture and R_s tend to be more sensitive to increased precipitation in more arid areas and more responsive to decreased precipitation in more humid areas. The responses of R_s and Q₁₀ were predominantly driven by precipitation‐induced changes in the soil moisture, whereas changes in the soil temperature had limited impacts. Finally, our synthesis of prolonged drought experiments also emphasizes the importance of the timing and frequency of precipitation events on ecosystem C cycles. Given these findings, we urge future studies to focus on manipulating the frequency, intensity, and seasonality of precipitation with an aim to improving our ability to predict and model feedback between R_s and climate change.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Precipitation variability does not affect soil respiration and nitrogen dynamics in the understorey of a Mediterranean oak woodland

Background and aims

Future climate scenarios for the Mediterranean imply increasing precipitation variability. This study presents a large-scale water manipulation experiment simulating changes in precipitation variability, aiming at a better understanding of the effects of rainfall patterns on soil C and N cycling and understorey productivity in a Mediterranean oak woodland.

Methods

We used rain-out shelters to achieve (1) a normal dry period (7 days), and (2) a dry period increased three-fold (21 days), without altering total annual precipitation inputs.

Results

The temporal patterns of soil respiration (R _s) and soil inorganic N were not affected by treatment. However, water infiltration and N leaching increased with large infrequent watering events. R _s and soil NH₄ ⁺-N correlated with soil temperature, with soil NO₃ ^?-N being influenced by leaching.

Conclusions

The lack of significant treatment effects on either R _s or soil inorganic N can be explained by (1) minor differences in plant productivity between the treatments, suggesting equal plant N demand, and (2) the absence of moisture dependence of R _s and soil NH₄ ⁺-N. Increased N leaching with large infrequent precipitation events may have longer-term consequences for ecosystem functioning. Our results contribute to an improved understanding of possible climate change effects on key ecosystem processes in Mediterranean ecosystems.     

Responses of soil respiration to soil management changes in an agropastoral ecotone in Inner Mongolia,China

Studying the responses of soil respiration (R_s) to soil management changes is critical for enhancing our understanding of the global carbon cycle and has practical implications for grassland management. Therefore, the objectives of this study were (1) quantify daily and seasonal patterns of R_s, (2) evaluate the influence of abiotic factors on R_s, and (3) detect the effects of soil management changes on R_s. We hypothesized that (1) most of daily and seasonal variation in R_s could be explained by soil temperature (T_s) and soil water content (S_w), (2) soil management changes could significantly affect R_s, and (3) soil management changes affected R_s via the significant change in abiotic and biotic factors. In situ R_s values were monitored in an agropastoral ecotone in Inner Mongolia, China, during the growing seasons in 2009 (August to October) and 2010 (May to October). The soil management changes sequences included free grazing grassland (FG), cropland (CL), grazing enclosure grassland (GE), and abandoned cultivated grassland (AC). During the growing season in 2010, cumulative R_s for FG, CL, GE, and AC averaged 265.97, 344.74, 236.70, and 226.42 gC m^?2 year^?1, respectively. The T_s and S_w significantly influenced R_s and explained 66%–86% of the variability in daily R_s. Monthly mean temperature and precipitation explained 78%–96% of the variability in monthly R_s. The results clearly showed that R_s was increased by 29% with the conversion of FG to CL and decreased by 35% and 11% with the conversion of CL to AC and FG to GE. The factors impacting the change in R_s under different soil management changes sequences varied. Our results confirm the tested hypotheses. The increase in Q₁₀ and litter biomass induced by conversion of FG to GE could lead to increased R_s if the climate warming. We suggest that after proper natural restoration period, grasslands should be utilized properly to decrease R_s.     

土壤碳氮特征沿干旱梯度时空格局及对气候变化的响应——基于中国生态系统研究网络(CERN)长期定位监测数据

土壤碳氮含量及其化学计量特征是表征生态系统碳汇能力和土壤质量的重要指标，在支撑生态系统结构功能以及缓解气候变化中起着关键作用。利用中国生态系统研究网络(CERN)长期定位监测数据，分析了土壤碳氮特征沿干旱梯度的时空规律及其对气候变化的响应。结果表明：空间上，典型荒漠草原生态系统随着干旱加剧，土壤有机碳和全氮含量减少，土壤有机碳对干旱响应的敏感性降低，而土壤全氮对干旱响应的敏感性增加，土壤有机碳随土壤全氮含量的增加而增加。时间上，2005—2018年，荒漠草原生态系统土壤有机碳和全氮含量变化速率沿干旱梯度表现出由负转正的增加趋势，其中，干旱区呈减少趋势，半干旱和半湿润地区呈增加趋势，鄂尔多斯站和沙坡头站呈显著增加趋势。从影响因素来看，土壤碳氮特征对降水量增加的敏感性沿干旱梯度呈现出先增强后减弱的“上凸”抛物线趋势，温度变化对土壤碳氮特征的调控没有表现出明显的干旱梯度效应。土壤碳氮比、土壤有机碳含量、土壤全氮含量对降水量和平均温度变化响应的敏感性均依次降低。不同干旱梯度土壤碳氮特征的变化规律为未来气候变化下生态系统结构与功能预测提供科学依据。     

Interannual variation in soil CO2 efflux and the response of root respiration to climate and canopy gas exchange in mature ponderosa pine

We examined a 6‐year record of automated chamber‐based soil CO₂ efflux (F_s) and the underlying processes in relation to climate and canopy gas exchange at an AmeriFlux site in a seasonally drought‐stressed pine forest. Interannual variability of F_s was large (CV=17%) with a range of 427 g C m^?2 yr^?1 around a mean annual F_s of 811 g C m^?2 yr^?1. On average, 76% of the variation of daily mean F_s could be quantified using an empirical model with year‐specific basal respiration rate that was a linear function of tree basal area increment (BAI) and modulated by a common response to soil temperature and moisture. Interannual variability in F_s could be attributed almost equally to interannual variability in BAI (a proxy for above‐ground productivity) and interannual variability in soil climate. Seasonal total F_s was twice as sensitive to soil moisture variability during the summer months compared with temperature variability during the same period and almost insensitive to the natural range of interannual variability in spring temperatures. A strong seasonality in both root respiration (R_r) and heterotrophic respiration (R_h) was observed with the fraction attributed to R_r steadily increasing from 18% in mid‐March to 50% in early June through early July before dropping rapidly to 10% of F_s by mid‐August. The seasonal pattern in R_r (10‐day averages) was strongly linearly correlated with tree transpiration (r²=0.90, P<0.01) as measured using sap flux techniques and gross ecosystem productivity (GEP, r²=0.83, P<0.01) measured by the eddy‐covariance approach. R_r increased by 0.43 g C m^?2 day^?1 for every 1 g C m^?2 day^?1 increase in GEP. The strong linear correlation of R_r to seasonal changes in GEP and transpiration combined with longer‐term interannual variability in the base rate of F_s, as a linear function of BAI (r²=0.64, P=0.06), provides compelling justification for including canopy processes in future models of F_s.     

Changes in biocrust cover drive carbon cycle responses to climate change in drylands

Dryland ecosystems account for ca. 27% of global soil organic carbon (C) reserves, yet it is largely unknown how climate change will impact C cycling and storage in these areas. In drylands, soil C concentrates at the surface, making it particularly sensitive to the activity of organisms inhabiting the soil uppermost levels, such as communities dominated by lichens, mosses, bacteria and fungi (biocrusts). We conducted a full factorial warming and rainfall exclusion experiment at two semiarid sites in Spain to show how an average increase of air temperature of 2–3 °C promoted a drastic reduction in biocrust cover (ca. 44% in 4 years). Warming significantly increased soil CO₂ efflux, and reduced soil net CO₂ uptake, in biocrust‐dominated microsites. Losses of biocrust cover with warming through time were paralleled by increases in recalcitrant C sources, such as aromatic compounds, and in the abundance of fungi relative to bacteria. The dramatic reduction in biocrust cover with warming will lessen the capacity of drylands to sequester atmospheric CO₂. This decrease may act synergistically with other warming‐induced effects, such as the increase in soil CO₂ efflux and the changes in microbial communities to alter C cycling in drylands, and to reduce soil C stocks in the mid to long term.     

Variation of morphological and chemical traits in sexes of the dioecious perennial grass Poa ligularis in relation to shrub cover and aridity in Patagonian ecosystems

Sexes of dioecious species may have dimorphic responses to environmental variation due to differences in resource requirements and reproductive costs. We analyzed the effect of aridity/relative shrub cover, and vicinity to shrub patches on morpho-chemical traits of sexes of the dioecious perennial grass Poa ligularis in patchy arid ecosystems in northern-central Patagonia. We hypothesized that sexes of P. ligularis have dimorphic responses in morpho-chemical traits in relation to the environmental variation induced by aridity/relative shrub cover and vicinity to shrub patches. We selected seven sites across a gradient of increasing aridity and relative shrub cover. We randomly collected 5–10 P. ligularis plants per site registering the sex (female or male) and location with respect to shrub patches (shrub patch or inter-patch). For each plant, we assessed morpho-chemical traits (height of the vegetative tillers, length/dry weight/area of blades, specific blade area, nitrogen and soluble phenol concentration in blades). Sexes showed dimorphic responses in height of vegetative tillers, blade length, and blade area with respect to vicinity to shrub patches; and in variation of soluble phenolics in blades in relation to aridity/relative shrub cover. Responses in both sexes were opposite to those expected by aridity, highlighting the role of favorable environments induced by shrub canopies on dimorphic responses of sexes of P. ligularis. Resource-rich microsites associated with shrub canopies promoted increased plant performance of females with high reproductive costs while resource-poor open areas, favorable for pollen dispersal, induced improved chemical defenses of males. These results are consistent with the resource availability hypothesis.     

Effects of grazing on grassland soil carbon: a global review

Soils of grasslands represent a large potential reservoir for storing CO₂, but this potential likely depends on how grasslands are managed for large mammal grazing. Previous studies found both strong positive and negative grazing effects on soil organic carbon (SOC) but explanations for this variation are poorly developed. Expanding on previous reviews, we performed a multifactorial meta‐analysis of grazer effects on SOC density on 47 independent experimental contrasts from 17 studies. We explicitly tested hypotheses that grazer effects would shift from negative to positive with decreasing precipitation, increasing fineness of soil texture, transition from dominant grass species with C₃ to C₄ photosynthesis, and decreasing grazing intensity, after controlling for study duration and sampling depth. The six variables of soil texture, precipitation, grass type, grazing intensity, study duration, and sampling depth explained 85% of a large variation (±150 g m^?2 yr^?1) in grazing effects, and the best model included significant interactions between precipitation and soil texture (P = 0.002), grass type, and grazing intensity (P = 0.012), and study duration and soil sampling depth (P = 0.020). Specifically, an increase in mean annual precipitation of 600 mm resulted in a 24% decrease in grazer effect size on finer textured soils, while on sandy soils the same increase in precipitation produced a 22% increase in grazer effect on SOC. Increasing grazing intensity increased SOC by 6–7% on C₄‐dominated and C₄–C₃ mixed grasslands, but decreased SOC by an average 18% in C₃‐dominated grasslands. We discovered these patterns despite a lack of studies in natural, wildlife‐dominated ecosystems, and tropical grasslands. Our results, which suggest a future focus on why C₃ vs. C₄‐dominated grasslands differ so strongly in their response of SOC to grazing, show that grazer effects on SOC are highly context‐specific and imply that grazers in different regions might be managed differently to help mitigate greenhouse gas emissions.     

Response of soil respiration to temperature and soil moisture: Effects of different vegetation types on a small scale in the eastern Loess Plateau of China

Soil respiration is affected by vegetation and environmental conditions. The purpose of this study was to investigate the effect of vegetation type on soil respiration, temperature and water content, and their correlations on a small scale. We measured soil respiration rate (R_s) over a 3-year period at biweekly intervals in three plots in the eastern Loess Plateau of China, with the same soil texture but different vegetation types: pine forest, grassland, and shrub land. Simultaneously, soil temperature (T_s) at 10 cm depth and soil water content (W_s) within 10 cm depth were measured. The seasonal course of R_s and T_s showed a similar temporal variation in the three plots, with higher values in summer and autumn and lower values in winter and spring. No significant differences (P>0.05) were found between plots, except for W_s. The mean cumulative release of CO₂ efflux from March to December was 962.5, 1027.5, and 1166.5 g C m^? 2 a^? 1 for plots 1, 2, and 3, respectively, with no significant difference between plots. The fitted exponential equations of R_s versus T_s from the 3-year data-set were significant (P < 0.05) with an R² of 0.72, 0.64, and 0.72 for plots 1, 2, and 3, respectively. The calculated Q₁₀ from the parameters of the fitted equation was 3.57, 3.52, and 3.61, and the R₁₀ was 2.36, 2.03, and 2.37 μmol CO₂ m^? 2 s^? 1 for plots 1, 2, and 3, respectively. Compared with the T_s, the correlations between R_s and W_s were not significant for the three plots. However, if the T_s was above 10°C, then their correlation was significant, and W_s had an impact on R_s. Four combined regression equations including two variables of T_s and W_s could be well established to model correlations between R_s and both T_s and W_s. Our study demonstrated that the exponential and power model fitted best and no significant different correlations of combined equations existed between the three plots. These results show that vegetation type had little impact on R_s, T_s, W_s, and their correlations, as well as on related parameters such as Q₁₀ and R₁₀. Therefore, while doing R_s research in a horizontal patchy vegetation conditions on a small area, the sampling location of measurements should focus on vertical dominant vegetation and ignore patch vegetation so as to reduce field work load.