草灌植被转变对草地生态系统及其水碳过程的影响研究进展

气候变暖和过度放牧的共同作用使全球草地出现明显的灌丛化现象,灌木去除是草地灌丛化控制的重要方式,识别这些草灌植被转变对生态系统、生态水文、土壤侵蚀和侵蚀碳流失的影响对草地可持续管理具有重要意义。综述了草地灌木入侵及其控制对植物群落和土壤功能(如土壤有机碳)的影响,以及这些草灌植被变化对生态水文、土壤侵蚀和土壤侵蚀碳流失等水碳耦合过程的影响机制。针对目前草地灌木入侵和去除对植物群落、植被格局、水土过程和功能影响研究的薄弱环节,对未来相关研究提出以下建议：(1)需深化草灌植被转变对碳、氮等生物地球化学循环的影响机制研究,(2)需重视核磁共振光谱、生物标志物、同位素等新技术和植被格局的指数与连通性等新方法在草灌植被转变的水、碳等生态效应研究中的应用,(3)需加强草灌植被格局和生态水文、土壤侵蚀与土壤侵蚀碳等水碳过程的多要素、多过程和多尺度的综合研究。本文旨在为灌丛化草地科学有效的生态恢复与多目标的土地利用管理提供理论支撑。     

Seasonal and Long-Term Changes to Active-Layer Temperatures after Tall Shrubland Expansion and Succession in Arctic Tundra

Vegetation, active-layer soils, and snow cover regulate energy exchange between the atmosphere and permafrost. Therefore, interactions between changes to tundra vegetation and soil thermal regime will fundamentally affect permafrost in a warmer world. We recorded soil temperatures for approximately 1 year in a Siberian Low Arctic landscape with a known history of alder (Alnus) shrub expansion on disturbed microsites in patterned ground. We recorded near-surface soil temperatures and measured physical properties of soils and vegetation on sorted-circle microsites in four stages of shrubland development: (1) tundra lacking tall shrubs; (2) shrub colonization zones; (3) mature shrublands; and (4) paludified, long-established shrublands with thick soil organic layers. Summer soil temperatures declined with increasing shrub cover and soil organic thickness; shrub colonization suppressed cryoturbation, facilitating the development of continuous vegetation and a surface organic mat on circles. Compared to open tundra, mature shrubs cooled soils by up to 9 °C during summer, but warmed soils by greater than 10 °C in winter presumably because they developed highly insulative snowpacks. Paludified shrublands had the coldest summer active layers, but winter soil temperatures were much colder than mature shrublands and were similar to earlier stages. Our results indicate that although tall shrub establishment dramatically warms winter soils within decades, much of this warming is transient at paludification-prone sites because the buildup of wet peat favors cooling in winter and the stature and snow-trapping capacity of shrubs diminish over time. In the ecosystem we studied, shrub expansion has contrasting effects on active-layer temperatures both seasonally and over longer timescales due to successional processes.     

Vegetation type determines heterotrophic respiration in subalpine Australian ecosystems

Soils are the largest store of carbon in the biosphere and cool‐cold climate ecosystems are notable for their carbon‐rich soils. Characterizing effects of future climates on soil‐stored C is critical to elucidating feedbacks to changes in the atmospheric pool of CO₂. Subalpine vegetation in south‐eastern Australia is characterized by changes over short distances (scales of tens to hundreds of metres) in community phenotype (woodland, shrubland, grassland) and in species composition. Despite common geology and only slight changes in landscape position, we measured striking differences in a range of soil properties and rates of respiration among three of the most common vegetation communities in subalpine Australian ecosystems. Rates of heterotrophic respiration in bulk soil were fastest in the woodland community with a shrub understorey, slowest in the grassland, and intermediate in woodland with grass understorey. Respiration rates in surface soils were 2.3 times those at depth in soils from woodland with shrub understorey. Surface soil respiration in woodlands with grass understorey and in grasslands was about 3.5 times that at greater depth. Both Arrhenius and simple exponential models fitted the data well. Temperature sensitivity (Q₁₀) varied and depended on the model used as well as community type and soil depth – highlighting difficulties associated with calculating and interpreting Q₁₀. Distributions of communities in these subalpine areas are dynamic and respond over relatively short time‐frames (decades) to changes in fire regime and, possibly, to changes in climate. Shifts in boundaries among communities and possible changes in species composition as a result of both direct and indirect (e.g. via fire regime) climatic effects will significantly alter rates of respiration through plant‐mediated changes in soil chemistry. Models of future carbon cycles need to take into account changes in soil chemistry and rates of respiration driven by changes in vegetation as well as those that are temperature‐ and moisture‐driven.     

Response of dominant grass and shrub species to water manipulation: an ecophysiological basis for shrub invasion in a Chihuahuan Desert Grassland

Increases in woody vegetation and declines in grasses in arid and semi-arid ecosystems have occurred globally since the 1800s, but the mechanisms driving this major land-cover change remain uncertain and controversial. Working in a shrub-encroached grassland in the northern Chihuahuan Desert where grasses and shrubs typically differ in leaf-level nitrogen allocation, photosynthetic pathway, and root distribution, we asked if differences in leaf-level ecophysiology could help explain shrub proliferation. We predicted that the relative performance of grasses and shrubs would vary with soil moisture due to the different morphological and physiological characteristics of the two life-forms. In a 2-year experiment with ambient, reduced, and enhanced precipitation during the monsoon season, respectively, the encroaching C(3) shrub (honey mesquite Prosopis glandulosa) consistently and substantially outperformed the historically dominant C(4) grass (black grama Bouteloua eriopoda) in terms of photosynthetic rates while also maintaining a more favorable leaf water status. These differences persisted across a wide range of soil moisture conditions, across which mesquite photosynthesis was decoupled from leaf water status and moisture in the upper 50 cm of the soil profile. Mesquite's ability to maintain physiologically active leaves for a greater fraction of the growing season than black grama potentially amplifies and extends the importance of physiological differences. These physiological and phenological differences may help account for grass displacement by shrubs in drylands. Furthermore, the greater sensitivity of the grass to low soil moisture suggests that grasslands may be increasingly susceptible to shrub encroachment in the face of the predicted increases in drought intensity and frequency in the desert of the southwestern USA.     

Dwarf shrub and grass vegetation resistant to long‐term experimental warming while microarthropod abundance declines on the Falkland Islands

Dwarf shrubs are a dominant plant type across many regions of the Earth and have hence a large impact on carbon and nutrient cycling rates. Climate change impacts on dwarf shrubs have been extensively studied in the Northern Hemisphere, and there appears to be large variability in response between ecosystem types and regions. In the Southern Hemisphere, less data are available despite dwarf shrub vegetation being a dominant feature of southern South America and mountainous regions of the Southern Hemisphere. Here, we present the response of an Empetrum rubrum dwarf shrub and a Poa grass community to 12 years of experimental climate manipulation achieved using open top chambers on the Falkland Islands, a cold temperate island group in the South Atlantic. The dwarf shrub and grass vegetation did not change significantly in cover, biomass or species richness over the 12 years period in response to climate warming scenarios of up to 1°C reflecting annual warming levels predicted in this region for the coming decades. The soil microarthropod community, however, responded with declines in abundance (37%) under warming conditions in the grass community, but no such changes were observed in the dwarf shrub community. Overall, our data indicate that dwarf shrub communities are resistant to the levels of climate warming predicted over the coming decades in the southern South America region and will, therefore, remain a dominant driver of local ecosystem properties.     

Pan-Arctic soil moisture control on tundra carbon sequestration and plant productivity

Long-term atmospheric CO₂ concentration records have suggested a reduction in the positive effect of warming on high-latitude carbon uptake since the 1990s. A variety of mechanisms have been proposed to explain the reduced net carbon sink of northern ecosystems with increased air temperature, including water stress on vegetation and increased respiration over recent decades. However, the lack of consistent long-term carbon flux and in situ soil moisture data has severely limited our ability to identify the mechanisms responsible for the recent reduced carbon sink strength. In this study, we used a record of nearly 100 site-years of eddy covariance data from 11 continuous permafrost tundra sites distributed across the circumpolar Arctic to test the temperature (expressed as growing degree days, GDD) responses of gross primary production (GPP), net ecosystem exchange (NEE), and ecosystem respiration (ER) at different periods of the summer (early, peak, and late summer) including dominant tundra vegetation classes (graminoids and mosses, and shrubs). We further tested GPP, NEE, and ER relationships with soil moisture and vapor pressure deficit to identify potential moisture limitations on plant productivity and net carbon exchange. Our results show a decrease in GPP with rising GDD during the peak summer (July) for both vegetation classes, and a significant relationship between the peak summer GPP and soil moisture after statistically controlling for GDD in a partial correlation analysis. These results suggest that tundra ecosystems might not benefit from increased temperature as much as suggested by several terrestrial biosphere models, if decreased soil moisture limits the peak summer plant productivity, reducing the ability of these ecosystems to sequester carbon during the summer.     

人工小叶锦鸡儿(Caragana microphylla)灌丛土壤水分动态研究

采用由植被空间序列断时间系列的方法。分析了1984,1987,1995和1999年建立的人工小叶锦鸡儿固沙植被土壤水分的时空变化特征,结果表明,受沙地土壤机械组成的影响,沙地土壤组成以物理性沙粒为主,＞0.01mm沙粒占97％以上,凋萎湿度为1．55％,田间持水量5．5%,大有效水为3．95％,随着固沙植被优势度的增加,小叶锦鸡儿灌丛下土壤含水量持续下降,1984年建立的植被区土壤含水量明显低于1999年建立的植被区;在年内土壤水分季节变化中,4个阶段植被区土壤水分状况在4－6月不断降低,到6月达到最低值,7－10月逐渐回升;对于建立较早的植被区（1984和1987年）,土壤水分垂直变化表现出随着土层深度的增加,土壤含水量逐递减的趋势,特别是70cm下土层中,土壤含水下降明显,含水量低于凋萎湿度,由于对根际区域土壤水分的利用,加剧了固沙植被区深层土壤（70cm下）水分的亏缺,进而影响植被物种的组成,随着小叶锦鸡儿灌丛年龄的增加,浅根性植物所占比重增加。     

Biological Soil Crust Microsites Are the Main Contributor to Soil Respiration in a Semiarid Ecosystem

Biological soil crusts (BSCs) are a key biotic component of dryland ecosystems worldwide. However, most studies carried out to date on carbon (C) fluxes in these ecosystems, such as soil respiration, have neglected them. We conducted a 3.5-year field experiment to evaluate the spatio-temporal heterogeneity of soil respiration in a semiarid Stipa tenacissima steppe and to assess the contribution of BSC-dominated areas to the annual soil respiration of the whole ecosystem. We selected the six most frequent microsites in the study area: Stipa tussocks (ST), Retama sphaerocarpa shrubs (RS), and open areas with very low (<5% BSC cover, BS), low, medium and high cover of well-developed BSCs. Soil respiration rates did not differ among BSC-dominated microsites but were significantly higher and lower than those found in BS and ST microsites, respectively. A model using soil temperature and soil moisture accounted for over 85% of the temporal variation in soil respiration throughout the studied period. Using this model, we estimated a range of 240.4–322.6 g C m⁻² y⁻¹ released by soil respiration at our study area. Vegetated (ST and RS) and BSC-dominated microsites accounted for 37 and 42% of this amount, respectively. Our results indicate that accounting for the spatial heterogeneity in soil respiration induced by BSCs is crucial to provide accurate estimations of this flux at the ecosystem level. They also highlight that BSC-dominated areas are the main contributor to the total C released by soil respiration and, therefore, must be considered when estimating C budgets in drylands.     

Spatially explicit patterns in a dryland's soil respiration and relationships with climate,whole plant photosynthesis and soil fertility

Arid and semiarid ecosystems play a significant role in regulating global carbon cycling, yet our understanding of the controls over the dominant pathways of dryland CO₂ exchange remains poor. Substantial amounts of dryland soil are not covered by vascular plants and this patchiness in cover has important implications for spatial patterns and controls of carbon cycling. Spatial variation in soil respiration has been attributed to variation in soil moisture, temperature, nutrients and rhizodeposition, while seasonal patterns have been attributed to changes in moisture, temperature and photosynthetic inputs belowground. To characterize how controls over respiration vary spatially and temporally in a dryland ecosystem and to concurrently explore multiple potential controls, we estimated whole plant net photosynthesis (A_net) and soil respiration at four distances from the plant base, as well as corresponding fine root biomass and soil carbon and nitrogen pools, four times during a growing season. To determine if the controls vary between different plant functional types for Colorado Plateau species, measurements were made on the C₄ shrub, Atriplex confertifolia, and C₃ grass, Achnatherum hymenoides. Soil respiration declined throughout the growing season and diminished with distance from the plant base, though variations in both were much smaller than expected. The strongest relationship was between soil respiration and soil moisture. Soil respiration was correlated with whole plant A_net, although the relationship varied between species and distance from plant base. In the especially dry year of this study we did not observe any consistent correlations between soil respiration and soil carbon or nitrogen pools. Our findings suggest that abiotic factors, especially soil moisture, strongly regulate the response of soil respiration to biotic factors and soil carbon and nitrogen pools in dryland communities and, at least in dry years, may override expected spatial and seasonal patterns.     

半干旱黄土区坡面尺度柠条生长状况及影响要素分析

以半干旱黄土丘陵区典型小流域坡面大规模人工种植柠条林为例,基于坡面不同部位柠条生长状况和生境条件调查,定量分析了地形变化、土壤水分及灌木密度对柠条生长的直接、间接影响及其贡献率。结果表明:(1)东坡大株柠条生长明显好于南坡,下坡位柠条生长状况略好于中上坡位,其他各坡位之间柠条生长状况差异较小;(2)大株柠条生长与浅层土壤水分有正相关关系,而与灌木密度和深层土壤水分则呈负相关关系;大株柠条灌木高度、灌木纵截面积和冠幅体积对浅层土壤水分的响应敏感,冠幅长度对坡向和坡位的响应较为敏感,冠幅宽度对灌木密度的响应较为敏感;(3)地形和土壤水分变化解释了59.9%的大株柠条生长变异,其中坡向、坡位和浅层土壤水分是影响大株柠条生长的主导环境因子,它们分别解释了21.1%、16.0%和13.1%的柠条生长变化。研究认为半干旱黄土区人工植被恢复既要重视空间布局,也要在后期实施必要的管理措施以维持人工林地的稳定性。     

The temperature responses of soil respiration in deserts: a seven desert synthesis

The temperature response of soil respiration in deserts is not well quantified. We evaluated the response of respiration to temperatures spanning 67°C from seven deserts across North America and Greenland. Deserts have similar respiration rates in dry soil at 20°C, and as expected, respiration rates are greater under wet conditions, rivaling rates observed for more mesic systems. However, deserts differ in their respiration rates under wet soil at 20°C and in the strength of the effect of current and antecedent soil moisture on the sensitivity and magnitude of respiration. Respiration increases with temperature below 30°C but declines for temperatures exceeding 35°C. Hot deserts have lower temperature sensitivity than cold deserts, and insensitive or negative temperature sensitivities were predicted under certain moisture conditions that differed among deserts. These results have implications for large-scale modeling efforts because we highlight the unique behavior of desert soil respiration relative to other systems. These behaviors include variable temperature responses and the importance of antecedent moisture conditions for soil respiration.     

Organic matter turnover in a sagebrush steppe landscape

Laboratory incubations of¹⁵N-amended soils from a sagebrush steppe in south-central Wyoming indicate that nutrient turnover and availability have complex patterns across the landscape and between microsites. Total and available N and P and microbial C and N were highest in topographic depressions characterized by tall shrub communities. Net and gross N mineralization rates and respiration were also highest in these areas, but microbial efficiencies expressing growth relative to respiration cost were highest in soils of exposed ridgetop sites (prostrate shrub communities). Similar patterns occurred between shrub and intershrub soils, with greater nutrient availability under shrubs, but lower microbial efficiencies under shrubs than between. Surface soils had higher soil nutrient pools and N mineralization rates than subsurface soils, but N and C turnover and microbial efficiencies were lower in those surface soils. All soils decreased in respiration, mineralization, and immobilization rates during the 30-day incubation period, apparently approaching a steady-state substrate use. Soil microbial activity of the high organic matter accumulation areas was apparently more limited by labile substrate.     

The response of native species to removal of invasive exotic grasses in a seasonally dry Hawaiian woodland

Abstract. Non-native perennial grasses form 30% of the live understory biomass in seasonally dry, submontane forests in Hawaii Volcanoes National Park, yet their effects on native species are unknown. We removed these grasses from plots of 20 m × 20 m in 1991 and maintained removal and control areas over the next three years. Two fast growing shrub species, Dodonaea viscosa and Osteomeles anthylidifolia, increased in size significantly more in removal areas than in controls. Individuals of the most abundant shrub species, Styphelia tameiameia showed no net growth response to grass removal. They did, however, change their architecture: many branches along the mid and upper sections of the main trunk died and a proliferation of new leaves and shoots occurred in the lower 40 cm of trunk. Basal diameter increase was very small in Metrosideros polymorpha, the dominant tree species in these sites. All species except Styphelia had significantly increased leaf tissue nitrogen in removal plots by 18 months after removal when compared to shrubs in control areas suggesting that removal plot shrubs had greater access to soil nitrogen. Available soil-N pools, which were generally higher in the removal plots, support this interpretation. Light levels near the soil surface were also higher where grasses were removed than where they were present which may have contributed to increased shrub growth. By contrast, soil moisture was consistently lower where grasses were removed than where they were still present. Shrub tissue carbon isotope values were consistent with the interpretation that shrubs in removal plots had less rather than more water available to them. Hence, the increased growth observed in removal plot shrubs could not be due to release from moisture competition. Lastly, our results showed that seedlings of all woody species except Metrosideros were significantly more abundant in removal plots at both one and three years after removal and initially high sapling mortality was balanced by high recruitment into the sapling class. We believe that over time this will result in increased densities of native shrubs if grasses are kept out. With the presence of grasses, shrub growth in these woodlands is reduced and biomass is shifting towards grasses.     

Consequences of anthropogenic disturbances on soil seed bank diversity and nurse shrub effect in a semiarid rangeland

What is the effect of common land use histories on the diversity, richness, spatial distribution and abundance of the soil seed bank (SSB)? Does the effect change between different microsites under shrub canopy? To address these questions we selected seven sites with different grazing and fire histories in the Patagonian Monte desert. We took soil samples in seven microsites at different distances of the trunk along a windward/leeward transect through shrub patches to estimate the SSB of perennial grasses, shrubs and annuals. Shrubs SSB was scarce. The nurse shrub effect on perennial grass SSB was evident at sites with a low disturbance intensity (as sites without continuous grazing), and higher on the leeward side under the shrub canopy. High disturbance intensity (such as permanent grazing) promoted a decrease in perennial grass SSB and an increase in annuals SBB (especially non-native). Land use histories related to fire showed a moderated disturbance response (medium values for both groups). Differences between land use histories varied depending on the windward/leeward microsite from which the SSB was analysed. Our results suggest a nurse effect of the shrub patches on the spatial distribution and abundance of the SSB, but this effect decrease under continuous grazing. Fire seems to have a positive effect on perennial grass SSB. Nevertheless, fire characteristics must be controlled. Moreover, perennial grass SSB was almost depleted under continuous grazing, driving the grazed system towards a vulnerable state; and annual non-native species take advantage of this disturbance and dominate the SSB.     

The Influence of Spatial Patterns of Soil Moisture on the Grass and Shrub Responses to a Summer Rainstorm in a Chihuahuan Desert Ecotone

The cycling of surface water, energy, nutrients, and carbon is different between semiarid grassland and shrubland ecosystems. Although differences are evident when grasslands are compared to shrublands, the processes that contribute to this transition are more challenging to document. We evaluate how surface redistribution of precipitation and plant responses to the resulting infiltration patterns could contribute to the changes that occur during the transition from grassland to shrubland. We measured soil water potential under grasses (Bouteloua eriopoda), shrubs (Larrea tridentata) and bare soil and changes in plant water relations and gas exchange following a 15 mm summer storm in the grassland–shrubland ecotone at the Sevilleta National Wildlife Refuge in central New Mexico USA. Following the storm, soil water potential (Ψ_s) increased to 30 cm depth beneath both grass and shrub canopies, with the greatest change observed in the top 15 cm of the soil. The increase in Ψ_s was greater beneath grass canopies than beneath shrub canopies. Ψ_s under bare soil increased only to 5 cm depth. The substantial redistribution of rainfall and different rooting depths of the vegetation resulted in high Ψ_s throughout most of the rooting volume of the grasses whereas soil moisture was unchanged throughout a large portion of the shrub rooting volume. Consistent with this pattern, predawn water potential (Ψ_PD) of grasses increased more than 5 MPa to greater than −1 MPa whereas Ψ_PD of shrubs increased to −2.5 MPa, a change of less than 2 MPa. Transpiration increased roughly linearly with Ψ_PD in both grasses and shrubs. In grasses, assimilation was strongly correlated with Ψ_PD whereas there was no relationship in shrubs where assimilation showed no significant response to the pulse of soil moisture following the storm. These data show that preferential redistribution of water to grass canopies enhances transpiration and assimilation by grasses following large summer storms. This process may inhibit shrubland expansion at the ecotone during periods without extreme drought.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

Soil–litter mixing and microbial activity mediate decomposition and soil aggregate formation in a sandy shrub-invaded Chihuahuan Desert grassland

Drylands account globally for 30% of terrestrial net primary production and 20% of soil organic carbon. Present ecosystem models under predict litter decay in drylands, limiting assessments of biogeochemical cycling at multiple scales. Overlooked decomposition drivers, such as soil–litter mixing (SLM), may account for part of this model-measurement disconnect. We documented SLM and decomposition in relation to the formation of soil-microbial films and microbial extracellular enzyme activity (EEA) in the North American Chihuahuan Desert by placing mesh bags containing shrub (Prosopis glandulosa) foliar litter on the soil surface within contrasting vegetation microsites. Mass loss (in terms of k, the decay constant) was best described by the degree of SLM and soil-microbial film cover. EEA was greatest during periods of rapid litter decomposition and associated SLM. Soil-microbial film cover on litter surfaces increased over time and was greater in bare ground microsites (50% litter surface area covered) compared to shrub and grass microsites (37 and 33% covered, respectively). Soil aggregates that formed in association with decomposing leaf material had organic C and N concentrations 1.5–2× that of local surface soils. Micrographs of soil aggregates revealed a strong biotic component in their structure, suggesting that microbial decomposition facilitates aggregate formation and their C and N content. Decomposition drivers in arid lands fall into two major categories, abiotic and biotic, and it is challenging to ascertain their relative importance. The temporal synchrony between surface litter mass loss, EEA, biotic film development, and aggregate formation observed in this study supports the hypothesis that SLM enhances decomposition on detached litter by promoting conditions favorable for microbial processes. Inclusion of interactions between SLM and biological drivers will improve the ability of ecosystem models to predict decomposition rates and dynamics in drylands.     

Fine-scale spatial heterogeneity and seed size determine early seedling survival in a desert perennial shrub (<Emphasis Type="Italic">Ericameria nauseosa:</Emphasis> Asteraceae)

In heterogenous landscapes, seeds settle in some microsites more readily than others, independently of whether those microsites are suitable for germination and subsequent growth, and survival of seedlings. Wild seeds and seedlings of Ericameria nauseosa var. oreophila in the sand-dune ecosystem near Mono Lake, California show both concordance and conflict in where seeds are retained and where seedlings flourish. Using a field experiment, we followed performance of seeds and seedlings (i.e., the rate of germination and successful emergence, initial seedling size, seedling growth and survival over first growing season), by planting seeds of known sizes in pre-identified microsites (windward and lee aspects of interspaces between shrubs, under shrub canopies, and under snag canopies, respectively). Heavier seeds were more likely to germinate and emerge successfully and resulted in initially larger seedlings than lighter seeds. These initially larger seedlings subsequently remained larger over the growing season and lived longer than initially smaller seedlings. Independent of seed and initial seedling size, seedlings that germinated under the canopy of adult shrubs grew larger and survived longer than seedlings that germinated in interspaces (open space with little or no vegetation) or under snags (dead adult shrubs). Seedlings grown under the canopy of adult shrubs experienced significantly less solar radiation and wind-deposited sand than seedlings grown in interspace or snag microsites. Sand burial more than 1 cm was lethal for seedlings. The two variables of seed size and microsite type had by far greater impact on eventual growth and survival of seedlings than did aspect, and the effects of the former two variables were independent of each other. This study amplifies the body of work on E. nauseosa at this site demonstrating that the earliest events in the life history of this long-lived perennial shrub appear to persist through time despite the harsh and variable environment.     

Competitive effects of shrubs and grasses in prairie

We investigated the relative contributions of size and growth form (biomass allocation) to competitive effects between grasses and shrubs in western Canada for two years. We measured the effects of grasses and shrubs on each other at the population level using removal experiments in natural vegetation. In prairie where shrub abundance was low, shrubs suppressed grasses as much as grasses suppressed shrubs, even though shrubs had six times more standing crop. In adjacent brush clumps, however, where shrub standing crop was 37 times grass standing crop, shrubs suppressed grasses strongly, whereas grasses did not suppress shrubs. Shrubs reduced available soil nitrogen more strongly than grasses did, but shrubs and grasses did not differ in their effects on light or soil water. On a per-gram basis, however, shrubs had smaller effects on light, nitrogen, and water consumption than grasses did. In spite of their smaller per-gram effects on resources, the secondary growth of shrubs allowed them to accumulate more mass and height, and to eventually displace grasses. During this process, competition between the woody and the herbaceous growth form changed from symmetric to asymmetric.     

Tree photosynthesis modulates soil respiration on a diurnal time scale

To estimate how tree photosynthesis modulates soil respiration, we simultaneously and continuously measured soil respiration and canopy photosynthesis over an oak‐grass savanna during the summer, when the annual grass between trees was dead. Soil respiration measured under a tree crown reflected the sum of rhizosphere respiration and heterotrophic respiration; soil respiration measured in an open area represented heterotrophic respiration. Soil respiration was measured using solid‐state CO₂ sensors buried in soils and the flux‐gradient method. Canopy photosynthesis was obtained from overstory and understory flux measurements using the eddy covariance method. We found that the diurnal pattern of soil respiration in the open was driven by soil temperature, while soil respiration under the tree was decoupled with soil temperature. Although soil moisture controlled the seasonal pattern of soil respiration, it did not influence the diurnal pattern of soil respiration. Soil respiration under the tree controlled by the root component was strongly correlated with tree photosynthesis, but with a time lag of 7–12 h. These results indicate that photosynthesis drives soil respiration in addition to soil temperature and moisture.