Comparisons of ectomycorrhizae on pinyon pines (Pinus edulis; Pinaceae) across extremes of soil type and herbivory

Using field and greenhouse studies, we examined the relationships among pinyon pines (Pinus edulis), their ectomycorrhizal mutualists, and their moth herbivores as a function of soil fertility. We studied two soil types—the ash and cinder soils of the San Francisco volcanic field and nearby sandy loam soils. In the field, pinyons growing in cinders suffered from reduced moisture, negative nitrogen mineralization rates, low phosphate levels, reduced growth, and high moth herbivory relative to pinyons growing in sandy loam. Pinyons growing in cinders also had twofold higher levels of ectomycorrhizal colonization than their noncinder counterparts. Similarly, in the greenhouse, seedlings grown in cinders had higher levels of ectomycorrhizal colonization and greater numbers of ectomycorrhizae than seedlings grown in sandy loam. Seedling shoot growth was significantly enhanced by ectomycorrhizae in both soils. These patterns support three conclusions. First, field and greenhouse studies demonstrated that trees growing in nutrient-poor soils had higher levels of ectomycorrhizal colonization than trees growing in better soils. Second, across soil types, variation in ectomycorrhizal colonization was better predicted by soil fertility than by herbivory. However, herbivory negatively affected ectomycorrhizae in the stressful cinder environment. Third, although mycorrhizae can be parasitic under some conditions, ectomycorrhizae had mutualistic impacts on pinyon seedlings across the environmental extremes we studied.     

GENETIC DIFFERENTIATION AND HETEROZYGOSITY IN PINYON PINE ASSOCIATED WITH RESISTANCE TO HERBIVORY AND ENVIRONMENTAL STRESS

Arizona's Sunset Crater began erupting in 1064 AD and for the next 200 years buried over 2,000 km² in ash, cinders, and lava. Soil analyses indicate that pinyon pines (Pinus edulis) currently colonizing the cinder fields are faced with a highly stressful environment. Many of these pinyons suffer chronic, intense insect herbivory that reduces plant growth and eliminates female cone production. In contrast, herbivory among pinyons growing in neighboring sandy-loam soils is minimal. Furthermore, numerous trees within the heavily infested cinder field population suffer relatively low herbivory and maintain normal growth and reproduction. We used four polymorphic enzymes to examine the relationship between herbivore attack, environmental stress and genotypes of the adjacent cinder field, and sandy-loam soil pinyon populations. Our results demonstrate that 1) resistant trees display significant genetic differences and are more heterozygous for two enzymes associated with herbivory than susceptible trees; and 2) the cinder-soil pinyons exhibit significant genetic differences and are more heterozygous for an enzyme associated with environmental stress than the neighboring sandy-loam soil pinyons. We conclude that heterozygosity of specific or closely linked loci may facilitate pinyon resistance to herbivory and environmental stress, and that strong selection across narrow geographic boundaries resulted in rapid genetic differentiation of pinyon populations.     

Increased moth herbivory associated with environmental stress of pinyon pine at local and regional levels

 Using 6 years of observational and experimental data, we examined the hypothesis that water and nutrient stress increase the susceptibility of pinyon pine (Pinus edulis) to the stem- and cone-boring moth (Dioryctria albovittella). At two geographic levels, a local scale of 550 km² and a regional scale of 10,000 km², moth herbivory was strongly correlated with an edaphic stress gradient. At a local scale, from the cinder soils of Sunset Crater to nearby sandy-loam soils, nine of ten soil macro- and micronutrients, and soil water content were lowest in cinder-dominated soils. Herbivore damage was six times greater on trees growing in the most water and nutrient deficient site at Sunset Crater compared to sites with well-developed soils. Percentage silt-clay content of soil, which was highly positively correlated with soil nutrient and soil moisture at a local scale, accounted for 56% of the variation in herbivory at a regional scale among 22 sites. Within and across sites, increased stem resin flow was positively associated with reduced moth attack. On the basis of moth distribution across a stress gradient, we predicted that pinyons growing in highly stressful environments would show increased resistance to herbivores if supplemented with water and/or nutrients. We conducted a 6-year experiment at a high-stress site where individual trees received water only, fertilizer only, and water + fertilizer. Relative to control trees, stem growth and resin flow increased in all three treatments, but only significantly in the water + fertilizer treatment. Although there was no significant difference in herbivore damage among these three treatments, there was an overall reduction in herbivore damage on all treatment trees combined, compared to control trees. This experiment suggests that release from stress leads to increased resistance to insect attack and is consistent with our observational data. While other studies have predicted that short-term stress will result in herbivore outbreaks, our studies extend this prediction to chronically stressed host populations. Finally, while flush-feeders are not predicted to respond positively to stressed host plants, we found a positive association between herbivore attack and stressed pinyon populations. Received: 25 December 1995 / Accepted: 15 August 1996     

Genetic variation associated with chronic water and nutrient stress in pinyon pine

We examined how genotypic variation at the glycerate dehydrogenase (GLY) locus in pinyon pine might be affected by environmental stress and herbivory. We compared GLY allelic and genotypic frequencies among mature and juvenile trees growing in stressful cinder soil and adjacent sandy-loam soil. While no association was found with herbivory, three lines of evidence support the hypothesis that GLY slow homozygotes (SS) are selected for under conditions of environmental stress. 1) We found no differences at the GLY locus between juvenile and mature trees growing in sandy-loam soil. 2) However, in the stressful cinder soil we found a lower frequency of SS homozygotes in juvenile trees than in mature trees, suggesting that selection has occurred in one generation. 3) Using biomass as a measure of fitness, SS homozygotes were superior to other GLY genotypes in cinder soil. Two lines of evidence were inconsistent with the hypothesis of selection for SS homozygotes in stressful environments. 1) SS homozygote frequency between habitats was not significantly different, suggesting that too few generations have passed for differentiation to occur, and/or insufficient selection pressure. 2) The low frequency of SS homozygote juvenile trees growing in cinder soil suggests that counterbalancing selection may prevent SS homozygotes from increasing in the population.     

Precipitation pulse use by an invasive woody legume: the role of soil texture and pulse size

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or “pulses”. The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (Ψ_pd), the isotopic abundance of deuterium in stem water (δD), the abundance of ¹³C in soluble leaf sugar (δ¹³C), and percent volumetric soil water content (θ_v) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between Ψ_pd and δ¹³C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting–drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.     

Soil respiration response to climate change in Pacific Northwest prairies is mediated by a regional Mediterranean climate gradient

Soil respiration is expected to increase with rising global temperatures but the degree of response may depend on soil moisture and other local factors. Experimental climate change studies from single sites cannot discern whether an observed response is site‐dependent or generalizable. To deconvolve site‐specific vs. regional climatic controls, we examined soil respiration for 18 months along a 520 km climate gradient in three Pacific Northwest, USA prairies that represents increasingly severe Mediterranean conditions from north to south. At each site we implemented a fully factorial combination of 2.5–3 °C warming and 20% added precipitation intensity. The response of soil respiration to warming was driven primarily by the latitudinal climate gradient and not site‐specific factors. Warming increased respiration at all sites during months when soil moisture was not limiting. However, these gains were offset by reductions in respiration during seasonal transitions and summer drought due to lengthened periods of soil moisture limitation. The degree of this offset varied along the north–south climate gradient such that in 2011 warming increased cumulative annual soil respiration 28.6% in the northern site, 13.5% in the central site, and not at all in the southern site. Precipitation also stimulated soil respiration more frequently in the south, consistent with an increased duration of moisture limitation. The best predictors of soil respiration in nonlinear models were the Normalized Difference Vegetation Index (NDVI), antecedent soil moisture, and temperature but these models provided biased results at high and low soil respiration. NDVI was an effective integrator of climate and site differences in plant productivity in terms of their combined effects on soil respiration. Our results suggest that soil moisture limitation can offset the effect of warming on soil respiration, and that greater growing‐season moisture limitation would constrain cumulative annual responses to warming.     

Growth response of subalpine fir (Abies lasiocarpa) to climate in the Olympic Mountains, Washington, USA

Growth response of subalpine fir (Abies lasiocarpa) to climate was studied across its local geographical and elevation range in the Olympic Mountains, Washington. A dendroecological analysis of subalpine fir across a range of elevations (1350-1850 m) and annual precipitation (125-350 cm y^?1), was used to compare environmental factors affecting growth. Climate-growth relationships were explored using Pearson product-moment correlation coefficients; partial correlation analysis was used to assess relationships among site chronologies and climatic variables. Radial growth is negatively correlated with winter precipitation at high elevation and wet sites, but not at low and middle elevation dry sites. Growth is positively correlated with current growing season temperature at all sites; however, growth is negatively correlated with previous year August temperature, indicating that climate affects growth in subsequent years. Positive correlations between growth and summer precipitation during the growing season at low and middle elevation dry sites suggest that soil moisture is partially limiting to growth on these sites. If the climate of the Pacific Northwest becomes warmer and drier, then subalpine fir growth may increase at high elevation and wet sites, but may decrease at lower elevation dry sites in the Olympic Mountains. However, the growth response of subalpine fir to potentially rapid climate change will not be uniform because subalpine fir grows over a wide range of topographic features, habitats, and local climates at different geographical scales. A comparison of growth response to current growing season temperature suggests that the temperature-related growth response of subalpine fir is not adequately described by the parabolic curve used in JABOWA-based models.     

Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Effects of vegetation change and soil disturbance on ectomycorrhizas ofBetula platyphylla var.japonica: A test using seedlings planted in soils taken from various sites

The occurrence and character of different types of ectomycorrhizas of birch seedlings were investigated in soils from three naturally regenerating birch stands: a forest site, a clear-cut site, and a site recently disturbed by plowing. Birch grown in soil from an evergreen broad-leaved forest without birch was also studied. The rate of ectomycorrhizal formation in the soil from the evergreen broad-leaved forest was lower than that in the soil from the other three sites. The ectomycorrhizal formation of seedlings grown in soil from the clear-cut and plowed sites were the same as or higher than that in soil from the birch forest site. The largest number of ectomycorrhizal types were formed in soil from the birch forest site. In the soil from the plowed site, only one type of ectomycorrhiza was formed, and it was different from the dominant type formed in soils from the birch forest site and the clear-cut site. The results of this investigation showed that equal levels of ectomycorrhizas were formed in soils from the different birch stands, but the types formed were different among those sites. It is likely that the different ectomycorrhizal fungi were better adapted to the soil conditions at each of those sites.     

Unexpected patterns of sensitivity to drought in three semi-arid grasslands

Global climate models forecast an increase in the frequency and intensity of extreme weather events, including severe droughts. Based on multi-year relationships between precipitation amount and aboveground annual net primary production (ANPP), semi-arid grasslands are projected to be among the most sensitive ecosystems to changes in precipitation. To assess sensitivity to drought, as well as variability within the shortgrass steppe biome, we imposed moderate and severe rainfall reductions for two growing seasons in three undisturbed grasslands that varied in soil type and climate. We predicted strong drought-induced reductions in ANPP at all sites and greater sensitivity to drought in sites with lower average precipitation, consistent with continental-scale patterns. Identical experimental infrastructure at each site reduced growing season rainfall events by 50 or 80%, and significantly reduced average soil moisture in both years (by 21 and 46% of control levels, respectively). Despite reductions in soil moisture, ANPP responses varied unexpectedly-from no reduction in ANPP to a 51% decrease. Although sensitivity to drought was highest in the semi-arid grassland with lowest mean annual precipitation, patterns in responses to drought across these grasslands were also strongly related to rainfall event size. When growing season rainfall patterns were dominated by many smaller events, ANPP was significantly reduced by drought but not when rainfall patterns were characterized by large rain events. This interaction between drought sensitivity and rainfall event size suggests that ANPP responses to future droughts may be reduced if growing season rainfall regimes also become more extreme.     

Soil Microbial Community Response to Drought and Precipitation Variability in the Chihuahuan Desert

Increases in the magnitude and variability of precipitation events have been predicted for the Chihuahuan Desert region of West Texas. As patterns of moisture inputs and amounts change, soil microbial communities will respond to these alterations in soil moisture windows. In this study, we examined the soil microbial community structure within three vegetation zones along the Pine Canyon Watershed, an elevation and vegetation gradient in Big Bend National Park, Chihuahuan Desert. Soil samples at each site were obtained in mid-winter (January) and in mid-summer (August) for 2 years to capture a component of the variability in soil temperature and moisture that can occur seasonally and between years along this watershed. Precipitation patterns and amounts differed substantially between years with a drought characterizing most of the second year. Soils were collected during the drought period and following a large rainfall event and compared to soil samples collected during a relatively average season. Structural changes within microbial community in response to site, season, and precipitation patterns were evaluated using fatty acid methyl ester (FAME) and polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) analyses. Fungal FAME amounts differed significantly across seasons and sites and greatly outweighed the quantity of bacterial and actinomycete FAME levels for all sites and seasons. The highest fungal FAME levels were obtained in the low desert scrub site and not from the high elevation oak–pine forests. Total bacterial and actinomycete FAME levels did not differ significantly across season and year within any of the three locations along the watershed. Total bacterial and actinomycete FAME levels in the low elevation desert-shrub and grassland sites were slightly higher in the winter than in the summer. Microbial community structure at the high elevation oak–pine forest site was strongly correlated with levels of NH₄ ⁺–N, % soil moisture, and amounts of soil organic matter irrespective of season. Microbial community structure at the low elevation desert scrub and sotol grasslands sites was most strongly related to soil pH with bacterial and actinobacterial FAME levels accounting for site differences along the gradient. DGGE band counts of amplified soil bacterial DNA were found to differ significantly across sites and season with the highest band counts found in the mid-elevation grassland site. The least number of bands was observed in the high elevation oak–pine forest following the large summer-rain event that occurred after a prolonged drought. Microbial responses to changes in precipitation frequency and amount due to climate change will differ among vegetation zones along this Chihuahuan Desert watershed gradient. Soil bacterial communities at the mid-elevation grasslands site are the most vulnerable to changes in precipitation frequency and timing, while fungal community structure is most vulnerable in the low desert scrub site. The differential susceptibility of the microbial communities to changes in precipitation amounts along the elevation gradient reflects the interactive effects of the soil moisture window duration following a precipitation event and differences in soil heat loads. Amounts and types of carbon inputs may not be as important in regulating microbial structure among vegetation zones within in an arid environment as is the seasonal pattern of soil moisture and the soil heat load profile that characterizes the location.     

Comparison of ectomycorrhizas of Quercus garryana (Fagaceae) on serpentine and non-serpentine soils in southwestern Oregon

The diversity of ectomycorrhizal communities associated with Quercus garryana on and off serpentine soils was compared and related to landscape-level diversity. Serpentine soils are high in magnesium, iron, and heavy metals and low in fertility. In plant communities on serpentine soils, a high proportion of flowering plant species are endemic. At three sites with paired serpentine and nonserpentine soils in southwestern Oregon, we sampled Q. garryana roots and categorized ectomycorrhizas by morphotyping and by restriction fragment length patterns. Ectomycorrhizas were abundant at all sites; no single fungal species dominated in the ectomycorrhizas. Of 74 fungal species characterized by morphotype and pattern of restriction fragment length polymorphisms, 46 occurred on serpentine soils, and 32 were unique to serpentine soil. These species are potentially endemic to serpentine soil. Similarities in species composition between paired serpentine and nonserpentine soils were not significantly lower than among three serpentine sites or among three nonserpentine sites. We conclude that mycorrhizal communities associated with oaks on serpentine soil do not differ in species richness or species evenness from those on neighboring nonserpentine soil.     

Trigonella arcuata-associated rhizobia��an Ensifer (Sinorhizobium) meliloti population adapted to a desert environment

Long-term monitoring of soil properties reveals site-specific ecosystem shifts in soil processes due to land use and climate changes. This paper aims to study the effects of physical landscape changes associated with grazing on soil thermal and moisture regime at the plot scale in a semiarid Leymus chinensis steppe of Inner Mongolia, China. The investigated sites were subjected to three grazing intensities: ungrazed since 1979 (UG79), moderately grazed only in winter time (WG), and heavily grazed (HG). At each plot, we recorded the soil moisture and temperature over a 6-year period that spanned between June 2004 and September 2009 and experienced a large range in precipitation (162 to 362 mm). Based on these monitoring data, we divided a year into four hydric periods: (1) growing period (late April to August); (2) transitional period from summer to winter (September?COctober); (3) winter time (November?Cfirst March); and (4) transitional period from winter to summer (March?CApril). In general, soil moisture in grazed sites was lower than in the ungrazed site, particularly for the 30?C50 cm soil layer. Seasonal fluctuation of the soil moisture, due to variable precipitation and atmospheric demands, was most significant in the topsoil (0?C10 cm) and was less pronounced in deeper soil. Regardless of hydric seasons, soil moisture was significantly influenced by grazing intensity, whereas soil temperature was slightly influenced. With increasing grazing intensity, soil water storage decreased remarkably. Consequently, grazing reduced plant available water and therefore grassland productivity, which are linked to a great extent with the trampling-induced soil structure change and soil moisture regime.     

全球变暖对太行山植被生产力及土壤水分的影响

将相同的自然植被用Lysimeter从高海拔下移至低海拔,温度升高2℃,同时设置平均降水,增加10％降水,增加20％降水,减少10％降水和减少20％降水5个降水处理,模拟全球变暖带来的温度升高和降水变化对植被生产力和土壤水分的影响,两年的野外实验表明,温度升高造成生态适应性差的野古草（Arundinelia hirta)生产力显著下降,致使整个测试群落生产力降低,低海拔实验点生产力显著低于高海拔实验点,温度升高对铁杆（Artemisia sacrorum)和黄背草（Themeda japonica)的影响较小,太行山区的植被对降水的变化反映,降水增加使植被的生产力水平显著提高,其中降水增加20％的处理生产力比平均降水处理的生产力增加22％,增加降水处理的土壤含水量显著高于平均降水和降水减少的实验处理,由于植被的减少,温度升高的实验点从第二年开始土壤水分较高。     

Tree-ring growth of Gmelin larch under contrasting local conditions in the north of Central Siberia

While the forest-tundra zone in Siberia, Russia has been dendroclimatologically well-studied in recent decades, much less emphasis has been given to a wide belt of northern taiga larch forests located to the south. In this study, climate and local site conditions are explored to trace their influence on radial growth of Gmelin larch (Larix gmelinii (Rupr.) Rupr.) trees developed on permafrost soils in the northern taiga. Three dendrochronological sites characterized by great differences in thermo-hydrological regime of soils were established along a short (ca. 100 m long) transect: on a river bank (RB), at riparian zone of a stream (RZ) and on a terrace (TER). Comparative analysis of the rate and year-to-year dynamics of tree radial growth among sites revealed considerable difference in both raw and standardized tree-ring width (TRW) chronologies obtained for the RZ site, characterized by shallow soil active layer depth and saturated soils. Results of dendroclimatic analysis indicated that tree-ring growth at all the sites is mostly defined by climatic conditions of a previous year and precipitation has stronger effect on TRW chronologies in comparison to the air temperatures. Remarkably, a great difference in the climatic response of TRW chronologies has been obtained for trees growing within a very short distance from each other. The positive relation of tree-ring growth with precipitation, and negative to temperature was observed in the dry site RB. In contrary, precipitation negatively and temperature positively influenced tree radial growth of larch at the water saturated RZ. Thus, a complicate response of northern Siberian larch forest productivity to the possible climate changes is expected due to great mosaic of site conditions and variability of environmental factors controlling tree-ring growth at different sites. Our study demonstrates the new possibilities for the future dendroclimatic research in the region, as various climatic parameters can be reconstructed from tree-ring chronologies obtained for different sites.     

Effect of temperature and precipitation on the annual diameter growth of Scots pine on drained peatlands and adjacent mineral soil sites in Finland

The climate conditions of the current and previous growing seasons have been shown to influence growth of coniferous trees in mineral soils sites. These dependencies may be different in peatlands where growth is generally more dependent on variations in soil water conditions. In the Nordic and Baltic countries, millions of hectares of peatlands and wetlands have been drained in order to enhance forest production. These drainage networks do not guarantee stable soil water conditions for the whole stand rotation. It is thus likely that precipitation in particular may have a different influence on annual growth in peatland to that in mineral soil sites. We studied the effect of precipitation and temperature on the inter-annual diameter growth of Scots pine (Pinus sylvestris L.) in Finland in drained peatland forests. The diameter growth data were limited to periods when growth response to drainage had levelled out. For comparison, growth data were also collected from adjacent mineral soil trees. The climate variables were monthly mean temperature and precipitation in a given location estimated from observations at the nearest weather stations by means of spatial smoothing. We used mixed linear models in describing the annual diameter growth of individual trees as a function of tree size and stand properties and expressed the residual variation as a function of climate parameters. The peatland and mineral soil growth variations showed different dependence on climate parameters. Peatland trees within 5 m of a ditch showed different climate responses compared to those located further away. Precipitation in July was negatively correlated with the diameter growth of peatland trees but there was no correlation with temperature. Growth of trees in mineral soils was positively correlated with March and April mean temperatures and May and June mean precipitation. The residual growth indices showed largely similar patterns in peatlands and mineral soil sites.     

Carbon dioxide exchange over multiple temporal scales in an arid shrub ecosystem near La Paz,Baja California Sur,Mexico

Arid environments represent 30% of the global terrestrial surface, but are largely under‐represented in studies of ecosystem carbon flux. Less than 2% of all FLUXNET eddy covariance sites exist in a hot desert climate. Long‐term datasets of these regions are vital for capturing the seasonal and interannual variability that occur due to episodic precipitation events and climate change, which drive fluctuations in soil moisture and temperature patterns. The objectives of this study were to determine the meteorological variables that drive carbon flux on diel, seasonal, and annual scales and to determine how precipitation events control annual net ecosystem exchange (NEE). Patterns of NEE from 2002 to 2008 were investigated, providing a record with multiple replicates of seasons and conditions. Precipitation was extremely variable (55–339 mm) during the study period, and reduced precipitation in later years (2004–2008) appears to have resulted in annual moderate to large carbon sources (62–258 g C m^?2 yr^?1) in contrast to the previously reported sink (2002–2003). Variations in photosynthetically active radiation were found to principally drive variations in carbon uptake during the wet growing season while increased soil temperatures at a 5 cm depth stimulated carbon loss during the dry dormant season. Monthly NEE was primarily driven by soil moisture at a 5 cm depth, and years with a higher magnitude of precipitation events showed a longer growing season with annual net carbon uptake, whereas years with lower magnitude had drier soils and displayed short growing seasons with annual net carbon loss. Increased precipitation frequency was associated with increased annual NEE, which may be a function of increased microbial respiration to more small precipitation events. Annual precipitation frequency and magnitude were found to have effects on the interannual variability of NEE for up to 2 years.     

Soil Respiration and Bacterial Structure and Function after 17 Years of a Reciprocal Soil Transplant Experiment

The effects of climate change on soil organic matter—its structure, microbial community, carbon storage, and respiration response—remain uncertain and widely debated. In addition, the effects of climate changes on ecosystem structure and function are often modulated or delayed, meaning that short-term experiments are not sufficient to characterize ecosystem responses. This study capitalized on a long-term reciprocal soil transplant experiment to examine the response of dryland soils to climate change. The two transplant sites were separated by 500 m of elevation on the same mountain slope in eastern Washington state, USA, and had similar plant species and soil types. We resampled the original 1994 soil transplants and controls, measuring CO₂ production, temperature response, enzyme activity, and bacterial community structure after 17 years. Over a laboratory incubation of 100 days, reciprocally transplanted soils respired roughly equal cumulative amounts of carbon as non-transplanted controls from the same site. Soils transplanted from the hot, dry, lower site to the cooler and wetter (difference of -5°C monthly maximum air temperature, +50 mm yr^-1 precipitation) upper site exhibited almost no respiratory response to temperature (Q₁₀ of 1.1), but soils originally from the upper, cooler site had generally higher respiration rates. The bacterial community structure of transplants did not differ significantly from that of untransplanted controls, however. Slight differences in local climate between the upper and lower Rattlesnake locations, simulated with environmental control chambers during the incubation, thus prompted significant differences in microbial activity, with no observed change to bacterial structure. These results support the idea that environmental shifts can influence soil C through metabolic changes, and suggest that microbial populations responsible for soil heterotrophic respiration may be constrained in surprising ways, even as shorter- and longer-term soil microbial dynamics may be significantly different under changing climate.     

Increased Soil Frost Versus Summer Drought as Drivers of Plant Biomass Responses to Reduced Precipitation: Results from a Globally Coordinated Field Experiment

Reduced precipitation treatments often are used in field experiments to explore the effects of drought on plant productivity and species composition. However, in seasonally snow-covered regions reduced precipitation also reduces snow cover, which can increase soil frost depth, decrease minimum soil temperatures and increase soil freeze–thaw cycles. Therefore, in addition to the effects of reduced precipitation on plants via drought, freezing damage to overwintering plant tissues at or below the soil surface could further affect plant productivity and relative species abundances during the growing season. We examined the effects of both reduced rainfall (via rain-out shelters) and reduced snow cover (via snow removal) at 13 sites globally (primarily grasslands) within the framework of the International Drought Experiment, a coordinated distributed experiment. Plant cover was estimated at the species level, and aboveground biomass was quantified at the functional group level. Among sites, we observed a negative correlation between the snow removal effect on minimum soil temperature and plant biomass production the next growing season. Three sites exhibited significant rain-out shelter effects on plant productivity, but there was no correlation among sites between the rain-out shelter effect on minimum soil moisture and plant biomass. There was no interaction between snow removal and rain-out shelters for plant biomass, although these two factors only exhibited significant effects simultaneously for a single site. Overall, our results reveal that reduced snowfall, when it decreases minimum soil temperatures, can be an important component of the total effect of reduced precipitation on plant productivity.     

Historical precipitation predictably alters the shape and magnitude of microbial functional response to soil moisture

Soil moisture constrains the activity of decomposer soil microorganisms, and in turn the rate at which soil carbon returns to the atmosphere. While increases in soil moisture are generally associated with increased microbial activity, historical climate may constrain current microbial responses to moisture. However, it is not known if variation in the shape and magnitude of microbial functional responses to soil moisture can be predicted from historical climate at regional scales. To address this problem, we measured soil enzyme activity at 12 sites across a broad climate gradient spanning 442–887 mm mean annual precipitation. Measurements were made eight times over 21 months to maximize sampling during different moisture conditions. We then fit saturating functions of enzyme activity to soil moisture and extracted half saturation and maximum activity parameter values from model fits. We found that 50% of the variation in maximum activity parameters across sites could be predicted by 30‐year mean annual precipitation, an indicator of historical climate, and that the effect is independent of variation in temperature, soil texture, or soil carbon concentration. Based on this finding, we suggest that variation in the shape and magnitude of soil microbial response to soil moisture due to historical climate may be remarkably predictable at regional scales, and this approach may extend to other systems. If historical contingencies on microbial activities prove to be persistent in the face of environmental change, this approach also provides a framework for incorporating historical climate effects into biogeochemical models simulating future global change scenarios.