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For many drylands, both long‐ and short‐term drought conditions can accentuate landscape heterogeneity at both temporal (e.g., role of seasonal patterns) and spatial (e.g., patchy plant cover) scales. Furthermore, short‐term drought conditions occurring over one season can exacerbate long‐term, multidecadal droughts or aridification, by limiting soil water recharge, decreasing plant growth, and altering biogeochemical cycles. Here, we examine how experimentally altered seasonal precipitation regimes in a mixed shrub grassland on the Colorado Plateau impact soil moisture, vegetation, and carbon and nitrogen cycling. The experiment was conducted from 2015 to 2019, during a regional multidecadal drought event, and consisted of three precipitation treatments, which were implemented with removable drought shelters intercepting ~66% of incoming precipitation including: control (ambient precipitation conditions, no shelter), warm season drought (sheltered April–October), and cool season drought (sheltered November–March). To track changes in vegetation, we measured biomass of the dominant shrub, Ephedra viridis, and estimated perennial plant and ground cover in the spring and the fall. Soil moisture dynamics suggested that warm season experimental drought had longer and more consistent drought legacy effects (occurring two out of the four drought cycles) than either cool season drought or ambient conditions, even during the driest years. We also found that E. viridis biomass remained consistent across treatments, while bunchgrass cover declined by 25% by 2019 across all treatments, with the earliest declines noticeable in the warm season drought plots. Extractable dissolved inorganic nitrogen and microbial biomass nitrogen concentrations appeared sensitive to seasonal drought conditions, with dissolved inorganic nitrogen increasing and microbial biomass nitrogen decreasing with reduced soil volumetric water content. Carbon stocks were not sensitive to drought but were greater under E. viridis patches. Additionally, we found that under E. viridis, there was a negative relationship between dissolved inorganic nitrogen and microbial biomass nitrogen, suggesting that drought‐induced increases in dissolved inorganic nitrogen may be due to declines in nitrogen uptake from microbes and plants alike. This work suggests that perennial grass plant–soil feedbacks are more vulnerable to both short‐term (seasonal) and long‐term (multiyear) drought events than shrubs, which can impact the future trajectory of dryland mixed shrub grassland ecosystems as drought frequency and intensity will likely continue to increase with ongoing climate change. [ABSTRACT FROM AUTHOR]

Mediterranean ecosystems are predicted to undergo longer and more intense summer droughts. The mechanisms underlying the response of herbaceous communities to such drier environments should be investigated to identify the resilience thresholds of Mediterranean rangelands. A 5‐year experiment was conducted in deep and shallow soil rangelands of southern France. A rainout shelter for 75 days in summer imposed drier and warmer conditions. Total soil water content was measured monthly to model available daily soil water. Aboveground net primary production (ANPP), forage quality, and the proportion of graminoids in ANPP were measured in spring and autumn. Plant senescence and plant cover were assessed in summer and spring, respectively. The experimental years were among the driest ever recorded at the site. Therefore, manipulated summer droughts were drier than long‐term ambient conditions. Interactions between treatment, community type, and experimental year were found for most variables. In shallow soil communities, spring plant cover decreased markedly with time. This legacy effect, driven by summer plant mortality and the loss of perennial graminoids, led to an abrupt loss of resilience when the extreme water stress index exceeded 37 mm 10 day−1, characterized by a reduction of spring plant cover below 50% and a decreased ANPP in rainy years. Conversely, the ANPP of deep soil communities remained unaffected by increased summer drought, although the presence of graminoids increased and forage nutritive value decreased. This study highlights the role of the soil water reserve of Mediterranean plant communities in modulating ecosystem responses to chronically intensified summer drought. Communities on deep soils were resilient, but communities on shallow soils showed a progressive, rapid, and intense degradation associated with a loss of resilience capacity. Notably, indexes of extreme stress were a better indicator of tipping points than indexes of integrated annual stress. Considering the role of soil water availability in other herbaceous ecosystems should improve the ability to predict the resilience of plant communities under climate change. [ABSTRACT FROM AUTHOR]

Human‐mediated dispersal of non‐native earthworms can cause substantial changes to the functioning and composition of ecosystems previously earthworm‐free. Some of these earthworm species have the potential to "geoengineer" soils and increase plant nitrogen (N) uptake. Yet the possible consequences of increased plant N concentrations on rodent grazing remains poorly understood. In this study, we present findings from a common garden experiment with two tundra communities, meadow (forb dominated) and heath (shrub dominated), half of them subjected to 4 years of earthworm presence (Lumbricus spp. and Aporrectodea spp.). Within four summers, our earthworm treatment changed plant community composition by increasing graminoid density by, on average, 94% in the heath vegetation and by 49% in the meadow. Rodent winter grazing was more intense on plants growing in soils with earthworms, an effect that coincided with higher N concentrations in plants, indicating a higher palatability. Even though earthworms reduced soil moisture, plant community productivity, as indicated by vegetation greenness (normalized difference vegetation index), was not negatively impacted. We conclude that earthworm‐induced changes in plant composition and trophic interactions may fundamentally alter the functioning of tundra ecosystems. [ABSTRACT FROM AUTHOR]

The potential effects of climate change on plant reproductive phenology include asynchronies with pollinators and reductions in plant fitness, leading to extinction and loss of ecosystem function. In particular, plant phenology is sensitive to extreme weather events, which are occurring with increasing severity and frequency in recent decades and are linked to anthropogenic climate change and shifts in atmospheric circulation. For 15 plant species in a Venezuelan cloud forest, we documented dramatic changes in monthly flower and fruit community composition over a 35‐year time series, from 1983 to 2017, and these changes were linked directly to higher temperatures, lower precipitation, and decreased soil water availability. The patterns documented here do not mirror trends in temperate zones but corroborate results from the Asian tropics. More intense droughts are predicted to occur in the region, which will cause dramatic changes in flower and fruit availability. [ABSTRACT FROM AUTHOR]

Tree roots not only acquire readily‐usable soil nutrients but also affect microbial decomposition and manipulate nutrient availability in their surrounding soils, that is, rhizosphere effects (REs). Thus, REs challenge the basic understanding of how plants adapt to the environment and co‐exist with other species. Yet, how REs vary among species in response to species‐specific bulk soil nutrient cycling is not well‐known. Here, we studied how plant‐controlled microbial decomposition activities in rhizosphere soils respond to those in their corresponding bulk soils and whether these relations depend on species‐specific nutrient cycling in the bulk soils. We targeted 55 woody species of different clades and mycorrhizal types in three contrasting biomes, namely a temperate forest, a subtropical forest, and a tropical forest. We found that microbial decomposition activities in rhizosphere soils responded linearly to those in their corresponding bulk soils at the species level. Thereafter, we found that REs (parameters in rhizosphere soils minus those in corresponding bulk soils) of microbial decomposition activities had negative linear correlations with microbial decomposition activities in corresponding bulk soils. A multiple factor analysis revealed that soil organic carbon, total nitrogen, and soil water content favored bulk soil decomposition activities in all three biomes, showing that the magnitude of REs varied along a fast‐slow nutrient cycling spectrum in bulk soils. The species of fast nutrient cycling in their bulk soils tended to have smaller or even negative REs. Therefore, woody plants commonly utilize both positive and negative REs as a nutrient‐acquisition strategy. Based on the trade‐offs between REs and other nutrient‐acquisition strategies, we proposed a push and pull conceptual model which can bring plant nutrient‐acquisition cost and plant carbon economics spectrum together in the future. This model will facilitate not only the carbon and nutrient cycling but also the mechanisms of species co‐existence in forest ecosystems. [ABSTRACT FROM AUTHOR]

Recurrent droughts are an inevitable consequence of climate change, yet how grasslands respond to such events is unclear. We conducted a 6‐year rainfall manipulation experiment in a semiarid grassland that consisted of an initial 2‐year drought (2015–2016), followed by a recovery period (2017–2018) and, finally, a second 2‐year drought (2019–2020). In each year, we estimated aboveground net primary productivity (ANPP), species richness, community‐weighted mean (CWM) plant traits, and several indices of functional diversity. The initial drought led to reduced ANPP, which was primarily driven by limited growth of forbs in the first year and grasses in the second year. Total ANPP completely recovered as the rapid recovery of grass productivity compensated for the slow recovery of forb productivity. The subsequent drought led to a greater reduction in total ANPP than the initial drought due to the greater decline of both grass and forb productivity. The structural equation models revealed that soil moisture influenced ANPP responses directly during the initial drought, and indirectly during the subsequent drought by lowering functional diversity, which resulted in reduced total ANPP. Additionally, ANPP was positively influenced by CWM plant height and leaf nitrogen during the recovery period and recurrent drought, respectively. Overall, the greater impact of the second drought on ecosystem function than the initial drought, as well as the underlying differential mechanism, underscores the need for an understanding of how increased drought frequency may alter semiarid grassland functioning. [ABSTRACT FROM AUTHOR]

Microorganisms can help plants and animals contend with abiotic stressors, but why they provide such benefits remains unclear. Here we investigated byproduct benefits, which occur when traits that increase the fitness of one species provide incidental benefits to another species with no direct cost to the provider. In a greenhouse experiment, microbial traits predicted plant responses to soil moisture such that bacteria with self‐beneficial traits in drought increased plant early growth, size at reproduction, and chlorophyll concentration under drought, while bacteria with self‐beneficial traits in well‐watered environments increased these same plant traits in well‐watered soils. Thus, microbial traits that promote microbial success in different moisture environments also promote plant success in these same environments. Our results demonstrate that byproduct benefits, a concept developed to explain the evolution of cooperation in pairwise mutualisms, can also extend to interactions between plants and nonsymbiotic soil microbes. [ABSTRACT FROM AUTHOR]

A foundational assumption of trait‐based ecology is that individual performances should be predicted by its functional traits. However, the trait–performance relationships reported in literature were typically weak, probably due to the ignorance of interactions between traits and environments, intraspecific trait variability, and hard traits (directly linked to physiological processes of interest). We conducted an experiment of planting 900 seedlings of six tree species separately (one seedling per pot) along experimentally manipulated light and water gradients, monitored their survival and growth, and measured their morphological, photosynthetic, and hydraulic traits. Most trait–performance relationships depended on the environments, either marginally changing (weak trait × environment interaction) or even reversing (strong trait × environment interaction) along light or water gradients in our experiment. Such trait × environment interactions were more likely to be detected in growth models using individual‐level traits than models using species mean traits, but seedling growth was not better modeled with individual‐level traits than species mean traits. Additionally, none of the hard traits (photosynthetic and hydraulic traits) were better predictors than soft traits (morphological traits) modeling seedling growth and survival along light and water gradients. Our study highlights the necessities of considering trait × environment interactions when predicting response of plants to changing environments. The benefits of using individual‐level traits or hard traits to predict plant performance might be reduced or even canceled if their measurement errors are not well controlled. [ABSTRACT FROM AUTHOR]

Mechanisms such as conspecific negative density dependence (CNDD) and niche partitioning have been proposed to explain species coexistence and community diversity. However, as a potentially important axis of niche partitioning, the role of interannual climate variability in driving local community dynamics remains largely unknown. Here we used a 15‐year monitoring data set of more than 53,000 seedlings in a temperate forest to examine (1) what are the relative effects of interannual climate variability, biotic interactions, and habitat conditions on seedling survival; (2) how the effects of biotic interactions change with interannual climate variability, and habitat conditions; and (3) whether the impacts of interannual climate variability, biotic interactions, and habitat conditions differ with plant traits. Interannual climate variability accounted for the most variation in seedling survival at the community level, followed by biotic interactions, and habitat conditions. Increased snowpack and decreased minimum temperature during the non‐growing season had positive effects on seedling survival. Effects of conspecific neighbor density were weakened in higher snowpack, effective accumulated temperature, elevation, and soil‐resource gradient, but were intensified with increased ultraviolet radiation, maximum precipitation, minimum temperature, and soil moisture. In addition, the relative importance of interannual climate variability versus biotic interactions differed depending on species‐trait groups. Specifically, biotic interactions for gravity‐dispersed species had a larger effect size in affecting seedling survival than other trait groups. Also, gravity‐dispersed species experienced a stronger CNDD than wind‐dispersed, probably because wind‐dispersed seedlings rarely had adult conspecifics nearby. We found that interannual climate variability was most strongly associated with seedling survival, but the magnitude of climatic effects varied among species‐trait groups. Interannual climate variability may act as an inhibitor or accelerator to density‐dependent interactions and should be accounted for in future studies, as both a potential direct and indirect factor in understanding the diversity of forest communities. [ABSTRACT FROM AUTHOR]

Understanding the consequences of altered rainfall patterns on litter decomposition is critical to predicting the feedback effect of climate change on atmospheric CO2 concentrations. Although their effect on microbial decomposition has received considerable attention, their effect on litter fragmentation by detritivores, the other dominant decomposition pathway, remains largely unexplored. Particularly, it remains unclear how different detritivore species and their interactions respond to changes in rainfall quantity and frequency. To fill this knowledge gap, we determined the contribution to litter decomposition of two detritivore species (millipede and isopod), separately and in combination, under contrasting rainfall quantity and frequency in a temperate forest. Although halving rainfall quantity and frequency decreased topsoil moisture by 7.8 and 13.1%, respectively, neither millipede‐ nor isopod‐driven decomposition were affected by these changes. In contrast, decomposition driven by both detritivore species in combination was 65.5% higher than expected based on monospecific treatments under high rainfall quantity, but unchanged or even lower under low rainfall quantity. This indicates that although detritivore activity is relatively insensitive to changes in rainfall patterns, large synergistic interactions between detritivore species may disappear under future rainfall patterns. Incorporating interspecific interactions between decomposers thus seems critical to evaluate the sensitivity of decomposition to altered rainfall patterns. [ABSTRACT FROM AUTHOR]

The species pool concept has advanced our understanding for how biodiversity is coupled at local and regional scales. However, it remains unclear how species pool size, the number of species available to disperse to a site, influences community assembly across spatial scales. We provide one of the first studies that assesses diversity across scales after experimentally assembling grassland communities from species pools of different sizes. We show that species pool size causes scale‐dependent effects on diversity in grasslands undergoing restoration by altering the shape of the species–area relationship (SAR). Specifically, larger species pools increased the slope of the SAR, but not the intercept, suggesting that dispersal from a larger pool causes species to be more spatially aggregated. This increased aggregation appears to be caused by sampling effects due to fewer individuals arriving per species, rather than stronger species sorting across variation in soil moisture. These scale‐dependent effects suggest that studies evaluating species pools at a single, small scale may underestimate their effects, thereby contributing to uncertainty about the importance of regional processes for community assembly and their consequences for ecological restoration. [ABSTRACT FROM AUTHOR]

As the atmosphere warms, precipitation events become larger, but less frequent. Yet, there is fundamental disagreement about how increased precipitation intensity will affect vegetation. Walter's two‐layer hypothesis and experiments testing it have demonstrated that precipitation intensity can increase woody plant growth. Observational studies have found the opposite pattern. Not only are the patterns contradictory, but inference is largely limited to grasslands and savannas. We tested the effects of increased precipitation intensity in a shrub‐steppe ecosystem that receives >30% of its precipitation as snow. We used 11 (8 × 8 m) shelters to collect and redeposit rain and snow as larger, more intense events. Total annual precipitation was the same in all plots, but each plot received different precipitation event sizes ranging from 1 to 18 mm. Over three growing seasons, larger precipitation event sizes increased soil water availability, sagebrush (Artemisia tridentata) stem radius, and canopy greenness, decreased new root growth in shallow soils, and had no effect on herbaceous plant cover. Thus, we found that increased precipitation intensity can increase soil water availability and woody plant growth in a cold semiarid system. Assuming that stem growth is positively correlated with shrub reproduction, establishment and spread, results suggest that increasing precipitation intensity may have contributed to the woody plant encroachment observed around the world in the past 50 yr. Further, continuing increases in precipitation intensity caused by atmospheric warming are likely to continue to contribute to shrub encroachment in the future. [ABSTRACT FROM AUTHOR]

Large‐scale warming will alter multiple local climate factors in alpine tundra, yet very few experimental studies examine the combined yet distinct influences of earlier snowmelt, higher temperatures and altered soil moisture on alpine ecosystems. This limits our ability to predict responses to climate change by plant species and communities. To address this gap, we used infrared heaters and manual watering in a fully factorial experiment to determine the relative importance of these climate factors on plant flowering phenology, and response differences among plant functional groups. Heating advanced snowmelt and flower initiation, but exposed plants to colder early‐spring conditions in the period prior to first flower, indicating that snowmelt timing, not temperature, advances flowering initiation in the alpine community. Flowering duration was largely conserved; heating did not extend average species flowering into the latter part of the growing season but instead flowering was completed earlier in heated plots. Although passive warming experiments have resulted in warming‐induced soil drying suggested to advance flower senescence, supplemental water did not counteract the average species advance in flowering senescence caused by heating or extend flowering in unheated plots, and variation in soil moisture had inconsistent effects on flowering periods. Functional groups differed in sensitivity to earlier snowmelt, with flower initiation most advanced for early‐season species and flowering duration lengthened only for graminoids and forbs. We conclude that earlier snowmelt, driven by increased radiative heating, is the most important factor altering alpine flowering phenology. Studies that only manipulate summer temperature will err in estimating the sensitivity of alpine flowering phenology to large‐scale warming. The wholesale advance in flowering phenology with earlier snowmelt suggests that alpine communities will track warming, but only alpine forbs and graminoids appear able to take advantage of an extended snow‐free season. [ABSTRACT FROM AUTHOR]

Grasses accumulate high concentrations of silicon (Si) in their tissues, with potential benefits including herbivore defense, improved water balance, and reduced leaf construction costs. Although Si is one of the most widely varying leaf constituents among individuals, species, and ecosystems, the environmental forces driving this variation remain elusive and understudied. To understand relationships between environmental factors and grass Si accumulation better, we analyzed foliar chemistry of grasses from 17 globally distributed sites where nutrient inputs and grazing were manipulated. These sites span natural gradients in temperature, precipitation, and underlying soil properties, which allowed us to assess the relative importance of soil moisture and nutrients across variation in climate. Foliar Si concentration did not respond to large mammalian grazer exclusion, but significant variation in herbivore abundance among sites may have precluded the observation of defoliation effects at these sites. However, nutrient addition consistently reduced leaf Si, especially at sites with low soil nitrogen prior to nutrient addition. Additionally, a leaf‐level trade‐off between Si and carbon (C) existed that was stronger at arid sites than mesic sites. Our results suggest soil nutrient limitation favors investment in Si over C‐based leaf construction, and that fixing C is especially costly relative to assimilating Si when water is limiting. Our results demonstrate the importance of soil nutrients and precipitation as key drivers of global grass silicification patterns. [ABSTRACT FROM AUTHOR]

Establishment and persistence are central to community assembly and are determined by how traits interact with the environment to determine performance (trait–environment interactions). Community assembly studies have rarely considered such trait–environment interactions, however, which can lead to incorrect inferences about how traits affect assembly. We evaluated how functional traits, environmental conditions, and trait–environment interactions structure plant establishment, as a measure of performance. Within 12 prairie restorations created by sowing 70 species, we quantified environmental conditions and counted individuals of each seeded species to quantify first‐year establishment. Three trait–environment interactions structured establishment. Leaf nitrogen interacted with herbivore pressure, as low leaf nitrogen species established relatively better under higher herbivory than species with high leaf nitrogen. Soil moisture interacted with root mass fraction (RMF), with low‐RMF species establishing better with low soil moisture and higher‐RMF species better on wetter soils. Specific leaf area (SLA) interacted with light availability, as low‐SLA species established better under high light conditions and high‐SLA species under low light conditions. Our work illustrates how community assembly can be better described by trait–environment interactions than correlating traits or environment with performance. This knowledge can assist species selection to maximize restoration success. [ABSTRACT FROM AUTHOR]

Recent studies have shown that invasive earthworms can dramatically reduce native biodiversity, both above and below the ground. However, we still lack a synthetic understanding of the underlying mechanisms behind these changes, such as whether earthworm effects on soil chemical properties drive such relationships. Here, we investigated the effects of invasive earthworms on soil chemical properties (pH, water content, and the stocks and fluxes of carbon, nitrogen, and phosphorus) by conducting a meta‐analysis. Invasive earthworms generally increased soil pH, indicating that the removal of organic layers and the upward transport of more base‐rich mineral soil caused a shift in soil pH. Moreover, earthworms significantly decreased soil water content, suggesting that the burrowing activities of earthworms may have increased water infiltration of and/or increased evapotranspiration from soil. Notably, invasive earthworms had opposing effects on organic and mineral soil for carbon and nitrogen stocks, with decreases in organic, and increases in mineral soil. Nitrogen fluxes were higher in mineral soil, whereas fluxes in organic soil were not significantly affected by the presence of invasive earthworms, indicating that earthworms mobilize and redistribute nutrients among soil layers and increase overall nitrogen loss from the soil. Invasive earthworm effects on element stocks increased with ecological group richness only in organic soil. Earthworms further decreased ammonium stocks with negligible effects on nitrate stocks in organic soil, whereas they increased nitrate stocks but not ammonium stocks in mineral soil. Notably, all of these results were consistent across forest and grassland ecosystems underlining the generality of our findings. However, we found some significant differences between studies that were conducted in the field (observational and experimental settings) and in the lab, such as that the effects on soil pH decreased from field to lab settings, calling for a careful interpretation of lab findings. Our meta‐analysis provides strong empirical evidence that earthworm invasion may lead to substantial changes in soil chemical properties and element cycling in soil. Furthermore, our results can help explain the dramatic effects of invasive earthworms on native biodiversity, for example, shifts towards the dominance of grass species over herbaceous ones, as shown by recent meta‐analyses. [ABSTRACT FROM AUTHOR]

Understanding the interactive effects of species invasions and climate change is essential for predicting future shifts in biodiversity. Because multiple stressors can interact in synergistic or antagonistic ways, it is notoriously difficult to anticipate their combined effects on species assemblages. However, some hypotheses predict that plant invasions will become increasingly problematic as climate change improves conditions for invaders or lowers the biotic resistance of native communities. In a 4‐yr field experiment, we quantified the individual and interactive effects of invasion by a globally problematic C4 grass, Imperata cylindrica, and chronic simulated drought imposed by rainout shelters on the whole plant communities of regenerating longleaf pine forest. Invasion both inhibited plant colonization and enhanced plot‐level extinctions, resulting in a severe (60%) loss of plant diversity across all functional groups, including perennial grasses and forbs, annual forbs, and woody species and dramatic shifts in community composition. Experimental drought reduced diversity by 20%, and caused a shift in the dominant functional groups, but had no significant effect on cover of the invader. The invader partially ameliorated water stress in the drought treatment such that invaded plots had higher soil moisture than uninvaded plots. Consequently, the combined effects of invasion and drought were lower than expected from an additive model of multiple stressors. These findings, which may have broader implications for how other C4 grass invaders will interact with drought to shift native community dynamics, challenge the perception that climate change will exacerbate invasions. In revealing that invasive species pose a major threat to the diversity and structure of native communities despite their moderating effects on abiotic stress, this work also highlights that management of aggressive invaders may be critical to preserving biodiversity regardless of future climate. [ABSTRACT FROM AUTHOR]

Intensifying drought is increasingly linked to global forest diebacks. Improved understanding of drought impacts on individual trees has provided limited insight into drought vulnerability in part because tree moisture access and depletion is difficult to quantify. In forests, moisture reservoir depletion occurs through water use by the trees themselves. Here, we show that drought impacts on tree fitness and demographic performance can be predicted by tracking the moisture reservoir available to trees as a mass balance, estimated in a hierarchical state‐space framework. We apply this model to multiple seasonal droughts with tree transpiration measurements to demonstrate how species and size differences modulate moisture availability across landscapes. The depletion of individual moisture reservoirs can be tracked over the course of droughts and linked to biomass growth and reproductive output. This mass balance approach can predict individual moisture deficit, tree demographic performance, and drought vulnerability throughout forest stands based on measurements from a sample of trees. [ABSTRACT FROM AUTHOR]

Abstract: Realistic representations of plant carbon exchange processes are necessary to reliably simulate biosphere–atmosphere feedbacks. These processes are known to vary over time and space, though the drivers of the underlying rates are still widely debated in the literature. Here, we measured leaf carbon exchange in >500 individuals of 98 species from the Neotropics to high boreal biomes to determine the drivers of photosynthetic and dark respiration capacity. Covariate abiotic (long‐ and short‐term climate) and biotic (plant type, plant size, ontogeny, water status) data were used to explore significant drivers of temperature‐standardized leaf carbon exchange rates. Using model selection, we found the previous week's temperature and soil moisture at the time of measurement to be a better predictor of photosynthetic capacity than long‐term climate, with the combination of high recent temperatures and low soil moisture tending to decrease photosynthetic capacity. Non‐trees (annual and perennials) tended to have greater photosynthetic capacity than trees, and, within trees, adults tended to have greater photosynthetic capacity than juveniles, possibly as a result of differences in light availability. Dark respiration capacity was less responsive to the assessed drivers than photosynthetic capacity, with rates best predicted by multi‐year average site temperature alone. Our results suggest that, across large spatial scales, photosynthetic capacity quickly adjusts to changing environmental conditions, namely light, temperature, and soil moisture. Respiratory capacity is more conservative and most responsive to longer‐term conditions. Our results provide a framework for incorporating these processes into large‐scale models and a data set to benchmark such models. [ABSTRACT FROM AUTHOR]

Abstract: Most forest ecosystems are simultaneously affected by concurrent global change drivers. However, when assessing these effects, studies have mainly focused on the responses to single factors and have rarely evaluated the joined effects of the multiple aspects of environmental change. Here, we analyzed the combined effects of anthropogenic nitrogen (N) deposition and climatic conditions on the radial growth of Acer saccharum, a dominant tree species in eastern North American forests. We capitalized on a long‐term N deposition study, replicated along a latitudinal gradient, that has been taking place for more than 20 yr. We analyzed tree radial growth as a function of anthropogenic N deposition (ambient and experimental addition) and of summer temperature and soil water conditions. Our results reveal that experimental N deposition enhances radial growth of this species, an effect that was accentuated as temperature increased and soil water became more limiting. The spatial and temporal extent of our data also allowed us to assert that the positive effects of growing under the experimental N deposition are likely due to changes in the physiological performance of this species, and not due to the positive correlation between soil N and soil water holding capacity, as has been previously speculated in other studies. Our simulations of tree growth under forecasted climate scenarios specific for this region also revealed that although anthropogenic N deposition may enhance tree growth under a large array of environmental conditions, it will not mitigate the expected effects of growing under the considerably drier conditions characteristic of our most extreme climatic scenario. [ABSTRACT FROM AUTHOR]

Abstract: Long‐term observations of ecological communities are necessary for generating and testing predictions of ecosystem responses to climate change. We investigated temporal trends and spatial patterns of soil fauna along similar environmental gradients in three sites of the McMurdo Dry Valleys, Antarctica, spanning two distinct climatic phases: a decadal cooling trend from the early 1990s through the austral summer of February 2001, followed by a shift to the current trend of warming summers and more frequent discrete warming events. After February 2001, we observed a decline in the dominant species (the nematode Scottnema lindsayae) and increased abundance and expanded distribution of less common taxa (rotifers, tardigrades, and other nematode species). Such diverging responses have resulted in slightly greater evenness and spatial homogeneity of taxa. However, total abundance of soil fauna appears to be declining, as positive trends of the less common species so far have not compensated for the declining numbers of the dominant species. Interannual variation in the proportion of juveniles in the dominant species was consistent across sites, whereas trends in abundance varied more. Structural equation modeling supports the hypothesis that the observed biological trends arose from dissimilar responses by dominant and less common species to pulses of water availability resulting from enhanced ice melt. No direct effects of mean summer temperature were found, but there is evidence of indirect effects via its weak but significant positive relationship with soil moisture. Our findings show that combining an understanding of species responses to environmental change with long‐term observations in the field can provide a context for validating and refining predictions of ecological trends in the abundance and diversity of soil fauna. [ABSTRACT FROM AUTHOR]

Anti-herbivore defense shows high levels of both inter- and intraspecific variability. Defending against herbivores may be costly to the plant when it requires a tradeoff in allocation between defense and other missed opportunities, such as reproduction. Indeed, the plastic expression of defensive traits allows the plant to invest resources in defense only when the risk of being damaged actually increases, avoiding wasted resources. Plants may assess risk by responding to volatile cues emitted by neighbors that are under attack. Most plastic responses likely depend on environmental conditions. In this experiment, we investigated the effect of water availability on resistance induced by volatile cues in sagebrush. We found that plants receiving additional water over summer and/or volatile cues from neighbor donor plants showed reduced herbivore damage compared to control plants. Interestingly, we found no evidence of interactions between additional water and volatile cues. We performed an inferential analysis comparing historical records of the levels of herbivore damage during different years that had different temperature and precipitation accumulations. Results confirmed findings from the experiment, as the regression model indicated that sagebrush was better defended during wetter and hotter seasons. Reports from the literature indicated that sagebrush is extremely sensitive to water availability in the soil. We suggest that water availability may directly affect resistance of herbivory as well as sensitivity to cues of damage. Costs and benefits of allocating resources to defensive traits may vary with environmental conditions. [ABSTRACT FROM AUTHOR]

Climate change is expected to alter primary production and community composition in alpine ecosystems, but the direction and magnitude of change is debated. Warmer, wetter growing seasons may increase productivity; however, in the absence of additional precipitation, increased temperatures may decrease soil moisture, thereby diminishing any positive effect of warming. Since plant species show individual responses to environmental change, responses may depend on community composition and vary across life form or functional groups. We warmed an alpine plant community at Niwot Ridge, Colorado continuously for four years to test whether warming increases or decreases productivity of life form groups and the whole community. We provided supplemental water to a subset of plots to alleviate the drying effect of warming. We measured annual above-ground productivity and soil temperature and moisture, from which we calculated soil degree days and adequate soil moisture days. Using an information-theoretic approach, we observed that positive productivity responses to warming at the community level occur only when warming is combined with supplemental watering; otherwise we observed decreased productivity. Watering also increased community productivity in the absence of warming. Forbs accounted for the majority of the productivity at the site and drove the contingent community response to warming, while cushions drove the generally positive response to watering and graminoids muted the community response. Warming advanced snowmelt and increased soil degree days, while watering increased adequate soil moisture days. Heated and watered plots had more adequate soil moisture days than heated plots. Overall, measured changes in soil temperature and moisture in response to treatments were consistent with expected productivity responses. We found that available soil moisture largely determines the responses of this forb-dominated alpine community to simulated climate warming. [ABSTRACT FROM AUTHOR]

Reciprocal transplant experiments have often provided evidence of local adaptation in temperate plants, but few such studies have been conducted in the tropics. To enhance our knowledge of local adaptation in tropical plants, we studied natural populations of two recently diverged Neotropical plant species, Costus allenii and C. villosissimus, in central Panama. We found that these species display a parapatric distribution that reflects local environmental differences on a fine geographic scale: C. allenii is found along ravines in the understory of primary forest, while C. villosissimus is found along forest edges. Light availability was lower in C. allenii habitats, while precipitation and soil moisture were lower in C. villosissimus habitats. We carried out reciprocal transplant experiments with seeds and clones of mature plants to test the hypothesis that the parapatric distribution of these species is due to divergent adaptation to their local habitats. We found strong evidence of local adaptation, i.e., when grown in their ''home'' sites, each species outperformed the species from an ''away'' site. Our finding that C. allenii and C. villosissimus are mainly isolated by their microhabitats provides a first step toward understanding the mechanisms of adaptation and speciation in the tropics. [ABSTRACT FROM AUTHOR]

Variation in mutualistic and antagonistic interactions are important sources of variation in population dynamics and natural selection. Environmental heterogeneity can influence the outcome of interactions by affecting the intensity of interactions, but also by affecting the demography of the populations involved. However, little is known about the relative importance of environmental effects on interaction intensities and demographic sensitivity for variation in population growth rates. We investigated how soil depth, soil moisture, soil nutrient composition, and vegetation height influenced the intensity of seed prédation as well as host plant demography and sensitivity to seed prédation in the perennial herb Primula farinosa. Intensity of seed prédation ranged from 0% to 80% of seeds damaged among the 24 study populations and was related to soil moisture in two of four years. The effect of seed prédation on plant population growth rate (λ) ranged from negligible to a reduction in X by 0.70. Sensitivity of population growth rate to prédation explained as much of the variation in the reductions in population growth rate due to seed prédation as did prédation intensity. Plant population growth rate in the absence of seed prédation and sensitivity to prédation were negatively related to soil depth and soil moisture. Both intensity of prédation and sensitivity to prédation were positively correlated with potential population growth rate and, as a result, there was no significant relationship between prédation intensity and realized population growth rate. We conclude that in our study system environmental context influences the effects of seed prédation on plant fitness and population dynamics in two important ways: through variation in interaction intensity and through sensitivity to the effects of this interaction. Moreover, our results show that a given abiotic factor can influence population growth rate in different directions through effects on potential growth rate, intensity of biotic interactions, and the sensitivity of population growth rate to interactions. [ABSTRACT FROM AUTHOR]

Changes in rainfall distribution, generally predicted by many climate models, can affect resource dynamics and ecosystem function. While little studied, intra-annual rainfall distribution may have particularly strong effects on competitive interactions. Here, we test whether increased rainfall event size and decreased frequency within a growing season can influence competitive dynamics related to the invasion of exotic annual grasses in California coastal sage scrub (CSS). We hypothesized that larger rainfall events and decreased frequency will increase the competitive ability of native CSS species: a deeper root system will permit greater water use during dry periods between pulses and enhance their resource depletion effect on more shallow-rooted grasses. We planted grass and CSS seedlings in an additive competition design under three rainfall treatments: frequent small events, infrequent large events, and infrequent small events. The first two treatments had the same total rainfall but different frequency, while the second and third treatments had the same frequency but different total rainfall. Rainfall treatment altered the competitive interactions between CSS and grasses. In the first year, the competitive effect of annual grasses on shrub seedlings was strongest under the frequent small rainfall regime where they reduced deep soil moisture and light. In year two, the established shrubs began to exert strong competitive effects on grasses, and these effects were strongest under the infrequent small rainfall regime (low total rain) where they reduced shallow soil moisture and decreased grass stomatal conductance. Results suggest that reductions in both rainfall frequency and total rainfall may be important to competitive interactions, and can alter plant community composition and invasion when species have different rooting depths and different responses to soil moisture. [ABSTRACT FROM AUTHOR]

Both biogeographical and rainfall manipulation studies show that soil water content can be a strong driver of microbial community composition. However, we do not yet know if these patterns emerge because certain bacterial taxa are better able to survive at dry soil moisture regimes or if they are due to other drought-sensitive ecosystem properties indirectly affecting microbial community composition. In this study, we evaluated (1) whether bacterial community composition changed under an 11-year drought manipulation and (2) whether shifts under drought could be explained by variation in the moisture sensitivity of growth among bacterial taxa (moisture niche partitioning). Using 454 pyrosequencing of 16S rRNA, we observed shifts in bacterial community composition under drought, coincident with changes in other soil properties. We wet-up dry soils from drought plots to five moisture levels, and measured respiration and the composition of actively growing communities using bromodeoxyuridine (BrdU) labeling of DNA. The field drought experiment affected the composition of the active community when incubated at different moisture levels in the laboratory, as well as short-term (36-hour) respiration rates. Independent of history, bacterial communities also displayed strong niche partitioning across the wet-up moisture gradient. Although this indicates that moisture has the potential to drive bacterial community composition under long-term drought, species distributions predicted by response to moisture did not reflect the community composition of plots that were subjected to long-term drought. Bacterial community structure was likely more strongly driven by other environmental factors that changed under long-term drought, or not shaped by response to water level upon wet-up. The approach that we present here for linking niches to community composition could be adapted for other environmental variables to aid in predicting microbial species distributions and community responses to environmental change. [ABSTRACT FROM AUTHOR]

Termites and large herbivores affect African savanna plant communities. Both functional groups are also important for nutrient redistribution across the landscape. We conducted an experiment to study how termites and large herbivores, alone and in combination, affect herbaceous species diversity patterns in an African savanna. Herbaceous vegetation on large vegetated Macrotermes mounds (with and without large herbivores) and on adjacent savanna areas (with and without large herbivores) was monitored over three years in Lake Mburo National Park, Uganda. We found substantial differences in species richness, alpha diversity, evenness, and stability between termite mound herbaceous vegetation and adjacent savanna vegetation. Within months of fencing, levels of species richness, evenness, and stability were no longer significantly different between savanna and mounds. However, fencing reduced the cumulative number of species, particularly for forbs, of which 48% of the species were lost. Fencing increased the beta diversity (dissimilarity among plots) on the resource-poor (in terms of both nutrients and soil moisture) savanna areas, while it did not significantly affect beta diversity on the resource-rich termite mounds. While termites cause substantial heterogeneity in savanna vegetation, large herbivores further amplify these differences by reducing beta diversity on the savanna areas. Large herbivores are, however, responsible for the maintenance of a large number of forbs at the landscape level. These findings suggest that the mechanisms underlying the effects of termites and large herbivores on savanna plant communities scale up to shape community structure and dynamics at a landscape level. [ABSTRACT FROM AUTHOR]

Soil microbes in temperate forest ecosystems are able to cycle several hundreds of kilograms of N per hectare per year and are therefore of paramount importance for N retention. Belowground C allocation by trees is an important driver of seasonal microbial dynamics and may thus directly affect N transformation processes over the course of the year. Our study aimed at unraveling plant controls on soil N cycling in a temperate beech forest at a high temporal resolution over a time period of two years, by investigating the effects of tree girdling on microbial N turnover. In both years of the experiment, we discovered (I) a summer N mineralization phase (between July and August) and (2) a winter N immobilization phase (November-February). The summer mineralization phase was characterized by a high N mineralization activity, low microbial N uptake, and a subsequent high N availability in the soil. During the autumn/winter N immobilization phase, gross N mineralization rates were low, and microbial N uptake exceeded microbial N mineralization, which led to high levels of N in the microbial biomass and low N availability in the soil. The observed immobilization phase during the winter may play a crucial role for ecosystem functioning, since it could protect dissolved N that is produced by autumn litter degradation from being lost from the ecosystem during the phase when plants are mostly inactive. The difference between microbial biomass N levels in winter and spring equals 38 kg N/ha and may thus account for almost one-third of the annual plant N demand. Tree girdling strongly affected annual N cycling: the winter N immobilization phase disappeared in girdled plots (microbial N uptake and microbial biomass N were significantly reduced, while the amount of available N in the soil solution was enhanced). This was correlated to a reduced fungal abundance in autumn in girdled plots. By releasing recently fixed photosynthates to the soil, plants may thus actively control the annual microbial N cycle. Tree belowground C allocation increases N accumulation in microorganisms during the winter which may ultimately feed back on plant N availability in the following growing season. [ABSTRACT FROM AUTHOR]

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Niño drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Niño drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems. [ABSTRACT FROM AUTHOR]

Two prominent mechanisms proposed to structure biodiversity are niche-based ecological filtering and chance arrival of propagules from the species pool. Seed arrival is hypothesized to play a particularly strong role in high-diversity plant communities with large potential species pools and many rare species, but few studies have explored how seed arrival and local ecological filters interactively assemble species-rich communities in space and time. We experimentally manipulated seed arrival and multiple ecological filters in high-diversity, herbaceous-dominated groundcover communities in longleaf pine savannas, which contain the highest small-scale species richness in North America (up to > 40 species/m²). We tested three hypotheses: (1) local communities constitute relatively open-membership assemblages, in which increased seed arrival from the species pool strongly increases species richness; (2) ecological filters imposed by local fire intensity and soil moisture influence recruitment and richness of immigrating species; and (3) ecological filters increase similarity in the composition of immigrating species. In a two-year factorial field experiment, we manipulated local fire intensity by increasing pre-fire fuel loads, soil moisture using rain shelters and irrigation, and seed arrival by adding seeds from the local species pool. Seed arrival increased species richness regardless of fire intensity and soil moisture but interacted with both ecological filters to influence community assembly. High-intensity fire decreased richness of resident species, suggesting an important abiotic filter. In contrast, high-intensity fire increased recruitment and richness of immigrating species, presumably by decreasing effects of other ecological filters (competition and resource limitation) in pos-tfire environments. Drought decreased recruitment and richness of immigrating species, whereas wet soil conditions increased recruitment but decreased or had little effect on richness. Moreover, some ecological filters (wet soil conditions and, to a lesser extent, high-intensity fire) increased similarity in the composition of immigrating species, illustrating conditions that influence deterministic community assembly in species-rich communities. Our experiment provides insights into how dispersal-assembly mechanisms may interact with niche-assembly mechanisms in space (spatial variation in disturbance) and time (temporal variation in resource availability) to structure high-diversity communities and can help guide conservation of threatened longleaf pine ecosystems in the face of habitat fragmentation and environmental change. [ABSTRACT FROM AUTHOR]

We provide an empirical, global test of the varying constraints hypothesis, which predicts systematic heterogeneity in the relative importance of biomass resources to burn and atmospheric conditions suitable to burning (weather/climate) across a spatial gradient of long-term resource availability. Analyses were based on relationships between monthly global wildfire activity, soil moisture, and mid-tropospheric circulation data from 2001 to 2007, synthesized across a gradient of long-term averages in resources (net primary productivity), annual temperature, and terrestrial biome. We demonstrate support for the varying constraints hypothesis, showing that, while key biophysical factors must coincide for wildfires to occur, the relative influence of resources to burn and moisture/weather conditions on fire activity shows predictable spatial patterns. In areas where resources are always available for burning during the fire season, such as subtropical/tropical biomes with mid-high annual long-term net primary productivity, fuel moisture conditions exert their strongest constraint on fire activity. In areas where resources are more limiting or variable, such as deserts, xeric shrublands, or grasslands/savannas, fuel moisture has a diminished constraint on wildfire, and metrics indicating availability of burnable fuels produced during the antecedent wet growing seasons reflect a more pronounced constraint on wildfire. This macro-scaled evidence for spatially varying constraints provides a synthesis with studies performed at local and regional scales, enhances our understanding of fire as a global process, and indicates how sensitivity to future changes in temperature and precipitation may differ across the world. [ABSTRACT FROM AUTHOR]

Climate models predict precipitation changes for much of the humid tropics, yet few studies have investigated the potential consequences of drought on soil carbon (C) cycling in this important biome. In wet tropical forests, drought could stimulate soil respiration via overall reductions in soil anoxia, but previous research suggests that litter decomposition is positively correlated with high rainfall fluxes that move large quantities of dissolved organic matter (DOM) from the litter layer to the soil surface. Thus, reduced rainfall could also limit C delivery to the soil surface, reducing respiration rates. We conducted a throughfall manipulation experiment to investigate how 25% and 50% reductions in rainfall altered both C movement into soils and the effects of those DOM fluxes on soil respiration rates. In response to the experimental drought, soil respiration rates increased in both the -25% and -50% treatments. Throughfall fluxes were reduced by 26% and 55% in the -25% and -50% treatments, respectively. However, total DOM fluxes leached from the litter did not vary between treatments, because the concentrations of leached DOM reaching the soil surface increased in response to the simulated drought. Annual DOM concentrations averaged 7.7 ± 0.8, 11.2 ± 0.9, and 15.8 ± 1.2mg C/L in the control, -25%, and -50% plots, respectively, and DOM concentrations were positively correlated with soil respiration rates. A laboratory incubation experiment confirmed the potential importance of DOM concentration on soil respiration rates, suggesting that this mechanism could contribute to the increase in CO2 fluxes observed in the reduced rainfall plots. Across all plots, the data suggested that soil CO2 fluxes were partially regulated by the magnitude and concentration of soluble C delivered to the soil, but also by soil moisture and soil oxygen availability. Together, our data suggest that declines in precipitation in tropical rain forests could drive higher CO2 fluxes to the atmosphere both via increased soil 02 availability and through responses to elevated DOM concentrations. [ABSTRACT FROM AUTHOR]

The dynamics of forest eëosystems depend largely on the survival of seedlings in their understories, but seedling survival is known to be limited by preemption of light and soil resources by overstory trees. It has been hypothesized that "common mycorrhizal networks," wherein roots of seedlings are linked to overstory trees by symbiotic mycorrhizal fungi, offset some or all of the negative effects of trees on seedlings. Here we report the results of an unambiguous experimental test of this hypothesis in a monodominant Pinus radiata forest. We also tested the hypothesis that adaptive differentiation among plant populations causes local plant genotypes to respond more positively to mycorrhizal networks than nonlocal plant genotypes. Our results demonstrate large positive effects of overstory mycorrhizal networks on seedling survival, along with simultaneous negative effects of tree roots, regardless of whether plant genotypes were locally derived. Physiological and leafchemistry measurements suggest that seedlings connected to common mycorrhizal networks benefited from increased access to soil water. The similar magnitude of the positive and negative overstory effects on seedlings and the ubiquity of mycorrhizal networks in forests suggest that mycorrhizal networks fundamentally influence the demographic and community dynamics of forest trees. [ABSTRACT FROM AUTHOR]

Competitive coexistence in a spatially heterogeneous environment is traditionally attributed to niche differences, but several recent theories have proposed important additional roles for propagule limitation and chance (e.g., neutral theory, stochastic niche theory, spatial storage effect). We tested whether propagule supply and timing of disturbance affected the coexistence of three ecologically similar plants that replace one another with partial overlap along a local soil gradient. We asked what prevents the species that dominates the most common habitat (Holocarpha virgata, open hillsides) from invading the habitats where the other two species are dominant (Calycadenia pauciflora, rocky hilltops; Hemizonia congesta, clay-rich bottomlands). We added abundant Holocarpha seeds into Calycadenia and Hemizonia habitats that were experimentally disturbed at different times of year. Initial Holocarpha seedling densities in Calycadenia and Hemizonia habitats equaled or exceeded those in unmanipulated Holocarpha habitat, but Holocarpha survival, adult size, and fecundity were much lower outside its own habitat. Holocarpha persisted in Calycadenia and Hemizonia habitats for three years, and springtime disturbance promoted this expansion. However, outside its own habitat Holocarpha showed below-replacement fitness and little competitive effect on the other two species. Our results were most consistent with a deterministic view of spatial niches. Nonetheless, chance events may often cause natural communities to include some transient populations at any given time, leading them to appear "unsaturated" with species. [ABSTRACT FROM AUTHOR]

The flowering of grasses is a process critical to plant population dynamics and genetics, herbivore performance, and human health. To better understand the climate factors governing grass flowering, we analyzed the patterns of culm production over 25 years for three perennial tallgrass prairie species at Konza Prairie in Kansas, USA. The three species (Andropogon gerardii, Sorghastrum nutans, and Schizachyrium scoparium) all utilize the C4 photosynthetic pathway and were measured annually at the same locations for the past 25 years in an annually burned watershed. Culm production of all three species increased with higher growing-season soil moisture and precipitation but differed in their responses to water availability at different times during the growing season. Relative to Andropogon, Sorghastrum responded more to precipitation early in the growing season, and Schizachyrium responded more to precipitation late in the growing season. Flowering by each species also revealed a threshold relationship with late-season soil moisture at ∼1 m depth, which likely is a proxy for season-long water balance. Although flowering can be influenced by conditions antecedent to the current growing season, neither soil moisture nor precipitation during the previous year influenced flowering over the 25-year period. Flowering culm production averaged 9% and 7% of total graminoid aboveground net primary production (ANPP) in the uplands and lowlands, respectively. Interannual variation in ANPP correlated only with Sorghastrum flowering, suggesting a predominant role of the species in ANPP responses to climate. [ABSTRACT FROM AUTHOR]

Large herbivores and topo-edaphic gradients are well-documented, major determinants of grassland plant production and species composition. In contrast, there is limited information about how these factors together may influence the composition of the arbuscular mycorrhizal fungus (AMF) communities associated with plants. AMF are a common component of grassland ecosystems where they can influence plant productivity, diversity, and soil stability. In this study, AMF community composition was analyzed in paired plots located inside and outside 40-44-year-old ungulate exclosures at six grassland sites in Yellowstone National Park (YNP), USA, that varied in soil moisture and the availability of soil nitrogen (N) and phosphorus (P). AMF spore abundance, species richness, and the relative abundance of AMF species were determined from soil samples collected (1) randomly (n = 5 samples) within each of the 12 plots and (2) from beneath the dominant grass (n = 5 samples per plot) at each site. Randomly collected soil samples explored the effects of ungulates and topographic position on AMF composition at the plant community level, subsuming potential effects of ungulates on plant species composition. Dominant plant samples examined how grazers, in particular, influenced AMF communities, while controlling for host-plant identity. Grazing decreased AMF spore abundance across the landscape (examined by random sampling) but increased the AMF species richness associated with dominant plants. Grazing influenced the AMF species composition at the plant community level and at the host-plant level by shifting the relative abundances of individual AMF species. Individual AMF species responded differently to grazing and N and P availability. Our results demonstrate how soil moisture and N and P availability across the landscape interact with grazing to influence AMF species composition. [ABSTRACT FROM AUTHOR]

Recent theory and research have highlighted how the brown (detritus-based) world may control the trophic structure of the green (grazing) world. Detritus can alter bottom-up control of green webs by affecting autotroph biomass and quality through its ability to alter ecosystem properties, including soil moisture and nutrient cycling. Additionally, the role of detritus as the food resource base of brown webs may subsidize omnivorous predators that can provide top-down control of green webs. Brown-green connections may be especially important following plant invasions, which often lead to increased detritus and altered food webs. I combine field experiments, observational data, and path analysis to understand how nonnative grasses impact native arthropod communities in a semiarid shrub system. Theory and correlative evidence predict that decreased shrub growth and nutritional quality, and increased feeding of detrital predators on the grazing web, would decrease the abundance of shrub arthropods. In contrast, I found nonnative litter increased shrub growth via increased soil moisture and produced a strong bottom-up increase of the grazing arthropod web; effects of detrital predators and plant quality were comparatively unimportant. I link these findings to the apparent lack of overlapping predators between the brown and green webs, and to the important abiotic role of litter in this xeric system, which increased native plants and the abundance and richness of arthropods on them. [ABSTRACT FROM AUTHOR]

The Canadian High Arctic has been warming for several decades. Over this period, tundra plant communities have been influenced by regional climate change, as well as other disturbances. At a site on Ellesmere Island, Nunavut, Canada, we measured biomass and composition changes in a heath community over 13 years using a point-intercept method in permanent plots (1995-2007) and over 27 years using a biomass harvest comparison (1981- 2008). Results from both methods indicate that the community became more productive over time, suggesting that this ecosystem is currently in transition. Bryophyte and evergreen shrub abundances increased, while deciduous shrub, forb, graminoid, and lichen cover did not change. Species diversity also remained unchanged. Because of the greater evergreen shrub cover, canopy height increased. From 1995 to 2007, mean annual temperature and growing season length increased at the site. Maximum thaw depth increased, while soil water content did not change. We attribute the increased productivity of this community to regional warming over the past 30-50 years. This study provides the first plot-based evidence for the recent pan-Arctic increase in tundra productivity detected by satellite-based remote-sensing and repeat-photography studies. These types of ground-level observations are critical tools for detecting and projecting long-term community-level responses to warming. [ABSTRACT FROM AUTHOR]

Abstract. The productivity of many tropical wet forests is generally limited by bioavailable phosphorus (P). Microbial activity is a key regulator of P availability in that it determines both the supply of P through organic matter decomposition and the depletion of bioavailable P through microbial uptake. Both microbial uptake and mineralization occur rapidly, and their net effect on P availability varies with soil moisture, temperature, and soil organic matter quantity and quality. Exploring the mechanisms driving P availability at fine temporal scales can provide insight into the coupling of carbon, water, and nutrient cycles, and ultimately, the response of tropical forests to climate change. Despite the recognized importance of P cycling to the dynamics of wet tropical forests and their potential sensitivity to short-term fluctuations in bioavailable P, the diurnal pattern of P remains poorly understood. This study quantifies diurnal fluctuations in labile soil P and evaluates the importance of biotic and abiotic factors in driving these patterns. To this end, measurements of labile P were made every other hour in a Costa Rican wet tropical forest oxisol. Spatial and temporal variation in Bray-extractable P were investigated in relation to ecosystem carbon flux, soil CO2 efflux, soil moisture, soil temperature, solar radiation, and sap-flow velocity. Spatially averaged bi-hourly (every two hours) labile P ranged from 0.88 to 2.48 μg/g across days. The amplitude in labile P throughout the day was 0.6 1-0.82 μg/g (4 1-54% of mean P concentrations) and was characterized by a bimodal pattern with a decrease at midday. Labile P increased with soil CO2 efflux and soil temperature and declined with increasing sap flow and solar radiation. Together, soil CO2 efflux, soil temperature, and sap flow explained 86% of variation in labile P. [ABSTRACT FROM AUTHOR]

A simple null model, particularly germane to small and vulnerable organisms such as parasites, is that local conditions set a stage upon which larger-scale dynamics play out. Soil moisture strongly influences survival of entomopathogenic nematodes (EPN), which in turn drive trophic cascades by protecting vegetation from root-feeding herbivores. In this study, we examine the mechanisms responsible for patchy occurrence of an entomopathogenic nematode, Hezerorhabditis mare/at us, in a California coastal prairie. One hypothesis proposes that biotic factors such as competition and natural enemies could regulate occurrence of EPN populations. We found that fungi and other enemies of EPN, although locally potent, did not explain the patterns of incidence across sites. Abiotic factors also have strong effects on EPN persistence, especially for vulnerable free-living stages. Thus, we tested the hypothesis that patchy occurrence of EPN on a large landscape was driven by differences in soil moisture. Our research uses long-term data on nematode incidence in combination with a landscape- level experiment to demonstrate the lack of a correlation between soil moisture and long-term persistence. A year-long experiment showed EPN mortality was weakly correlated with soil moisture among our study sites. Thirteen years of data, however, showed that colonization rates were highly correlated with long-term persistence. Sites with highest long-term persistence experienced the highest rates of rhizosphere colonization, extinction, and turnover. As a result, we concluded that metapopulation dynamics override limitations set by local and short-term abiotic conditions to determine long-term persistence in this parasite-driven trophic cascade. [ABSTRACT FROM AUTHOR]

Spatially periodic vegetation patterns, forming gaps, bands, labyrinths, or spots, are characteristic of arid and semiarid landscapes. Self-organization models can explain this variety of structures within a unified conceptual framework. All these models are based on the interplay of positive and negative effects of plants on soil water, but they can be divided according to whether they assume the interactions to be mediated by water redistribution through runoff/diffusion or by plants' organs. We carried out a multi-proxy approach of the processes operating in a gapped pattern in southwest Niger dominated by a shrub species. Soil moisture within the root layer was monitored in time and space over one month of the rainy season. Soil water recharge displayed no spatial variation with respect to vegetation cover, but the stock half-life under cover was twice that of bare areas. A kernel of facilitation by the aboveground parts of shrubs was parameterized, and soil water half-life was significantly correlated to the cumulated facilitative effects of shrubs. The kernel range was found to be smaller than the canopy radius (81%). This effect of plants on soil water dynamics, probably through a reduction of evaporation by shading, is shown to be a better explanatory variable than potentially relevant soil and topography parameters. The root systems of five individuals of Combreturn micranthum G. Don were excavated. Root density data were used as a proxy to parameterize a kernel function of interplant competition. The range of this kernel was larger than the canopy radius (125%). The facilitation-to-competition range ratio, reflecting the above-to-belowground ratio of plant lateral extent, was smaller than 1 (0.64), a result supporting models assuming that patterning may emerge from an adaptation of plant morphology to aridity and shallow soils by means of an extended lateral root system. Moreover, observed soil water gradients had directions opposite to those assumed by alternative mathematical models based on underground water diffusion. This study contributes to the growing awareness that combined facilitative and competitive plant interactions can induce landscape-scale patterns and shape the two-way feedback loops between environment and vegetation. [ABSTRACT FROM AUTHOR]

Hydrologic flow and connectivity act as important determinants of ecological pattern and process in heterogeneous landscapes. Here we examine how the routing of water through the drainage network of an upper Sonoran Desert basin influences landscape patterns of soil respiration (SR) at both seasonal and event-based timescales. At seasonal timescales, SR varied up to 13-fold with downstream position in the drainage network, and annual estimates of CO2 efflux ranged from 185 g C·m-2·yr-1 to 1190 g C·m-2·yr-1 for sites arrayed along the same flow path. Spatial patterns of SR were unrelated to the carbon and water content of surface soils, but rather tracked changes in plant size and productivity, which in turn reflect downstream increases in groundwater availability. The relative importance of precipitation and temperature as drivers of SR also changed with landscape position, with the latter becoming more important in downstream locations. At the scale of individual precipitation events, SR increased up to 30-fold upon rewetting but typically returned to background levels within 24 h, even when soil moisture remained elevated. Unlike patterns observed at seasonal scales, event-based losses of CO2 varied across the landscape as a function of the organic-matter content in surface soils. Results from labile carbon amendments confirm that CO2 losses following precipitation pulses are initially constrained by substrate availability, not soil drying. By mediating spatial patterns of vegetation structure and soil resource availability, drainage networks represent an important physical template upon which belowground processes are organized in desert basins. [ABSTRACT FROM AUTHOR]

A large proportion of rainfall in dryland ecosystems is intercepted by plant foliage and is generally assumed to evaporate to the atmosphere or drip onto the soil surface without being absorbed. We demonstrate foliar absorption of intercepted rainfall in a widely distributed, continental dryland, woody-plant genus: Juniperus. We observed substantial improvement in plant water status, exceeding 1.0 MPa water potential for drought-stressed plants, following precipitation on an experimental plot that excluded soil water infiltration. Experiments that wetted shoots with unlabeled and with isotopically labeled water confirmed that water potential responded substantially to foliar wetting, that these responses were not attributable to re-equilibration with other portions of the xylem, and that magnitude of response increased with water stress. Foliar absorption is not included in most ecological, hydrological, and atmospheric models; has implications for interpreting plant isotopic signatures; and not only supplements water acquisition associated with increases in soil moisture that follow large or repeated precipitation events, but also enables plants to bypass soil water uptake and benefit from the majority of precipitation events, which wet foliage but do not increase soil moisture substantially. Foliar absorption of intercepted water could be more important than previously appreciated, especially during drought when water stress is greatest. [ABSTRACT FROM AUTHOR]

A 10-year experiment tested for variation in competition intensity over time in a natural grassland at the northern edge of the Great Plains. Growing-season precipitation varied fivefold during the study. All ecosystem-level variables varied significantly among years, and most covaried in expected ways. The covers of all common grasses possessing the C3 photosynthetic pathway varied significantly among years; in contrast, all common species with traits associated with drought tolerance (a C4 grass, a lichen, a spikemoss, and a subshrub) did not vary. Annual transplant experiments measured the competitive effects of neighbors on the growth of individuals of the native grass Bouteloua gracilis. A significant interaction between year and competition showed that competition intensity varied among years. The size of this effect, however, was small (η2 = 0.074) relative to the size of the direct effect of competition (η2 = 0.20) or the year in which the experiment was conducted (η2 = 0.51). Further, competition intensity was not significantly related to any variable describing standing crop or resources, or species richness. Species richness was highest in years with high precipitation, standing crop, and individual growth, due to the recruitment of rare species that were absent from dry years. In summary, variation in competition intensity was statistically significant but had small effects relative to the direct effects of climate. [ABSTRACT FROM AUTHOR]

Human modification of the environment is causing both loss of species and changes in resource availability. While studies have examined how species loss at the local level can influence invasion resistance, interactions between species loss and other components of environmental change remain poorly studied. In particular, the manner in which native diversity interacts with resource availability to influence invasion resistance is not well understood. We created experimental plant assemblages that varied in native species (1-16 species) and/or functional richness (defined by rooting morphology and phenology; one to five functional groups). We crossed these diversity treatments with resource (water) addition to determine their interactive effects on invasion resistance to spotted knapweed (Centaurea maculosa), a potent exotic invader in the intermountain West of the United States. We also determined how native diversity and resource addition influenced plant-available soil nitrogen, soil moisture, and light. Assemblages with lower species and functional diversity were more heavily invaded than assemblages with greater species and functional diversity. In uninvaded assemblages, experimental addition of water increased soil moisture and plant-available nitrogen and decreased light availability. The availability of these resources generally declined with increasing native plant diversity. Although water addition increased susceptibility to invasion, it did not fundamentally change the negative relationship between diversity and invasibility. Thus, native diversity provided strong invasion resistance even under high resource availability. These results suggest that the effects of local diversity can remain robust despite enhanced resource levels that are predicted under scenarios of global change. [ABSTRACT FROM AUTHOR]

Climate warming-induced plant species loss is likely to be nonrandom and based on species-specific susceptibility to changing climate. We examined the ecological consequences of losing shallow-rooted forbs, a group of species we predict to be adversely affected by climate change based on their response to experimental warming. After three years of experimental species removal, tap-rooted forbs and grasses were able to fully compensate for the loss of shallow-rooted forbs with increased biomass production. Moreover, the remaining plant community yielded a larger biomass response to nitrogen addition when shallow-rooted forbs were removed, possibly because removal led to increased soil moisture. We conclude that, although shallow-rooted forbs share a common response to warming, their loss did not affect community-level biomass. However, the loss of shallow-rooted forbs could result in increased sensitivity to perturbations, such as changing nutrient availability. Our results demonstrate that realistic, nonrandom scenarios of species loss do not necessarily follow the general pattern of decreased productivity and dampened response to nitrogen addition with species loss that is predicted by theory and many experimental results. Further examinations of nonrandom species loss in other ecosystems are needed to further improve our understanding of the consequences of human-driven species loss. [ABSTRACT FROM AUTHOR]

Lianas (woody vines) are an important and dynamic component of many forests throughout the world, and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests. In this study, we used a 45-year data set to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin (USA) since 1959. We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge. We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years. However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges. Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies. [ABSTRACT FROM AUTHOR]