Coordination of leaf, root, and seed traits shows the importance of whole plant economics in two semiarid grasslands.

Uncertainty persists within trait-based ecology, partly because few studies assess multiple axes of functional variation and their effect on plant performance. For 55 species from two semiarid grasslands, we quantified: (1) covariation between economic traits of leaves and absorptive roots, (2) covariation among economic traits, plant height, leaf size, and seed mass, and (3) relationships between these traits and species' abundance. Pairs of analogous leaf and root traits were at least weakly positively correlated (e.g. specific leaf area (SLA) and specific root length (SRL)). Two pairs of such traits, N content and DMC of leaves and roots, were at least moderately correlated (r > 0.5) whether species were grouped by site, taxonomic group and growth form, or life history. Root diameter was positively correlated with seed mass for all groups of species except annuals and monocots. Species with higher leaf dry matter content (LDMC) tended to be more abundant (r = 0.63). Annuals with larger seeds were more abundant (r = 0.69). Compared with global-scale syntheses with many observations from mesic ecosystems, we observed stronger correlations between analogous leaf and root traits, weaker correlations between SLA and leaf N, and stronger correlations between SRL and root N. In dry grasslands, plant persistence may require coordination of above- and belowground traits, and dense tissues may facilitate dominance.

Shower thoughts: why scientists should spend more time in the rain.

Stormwater is a vital resource and dynamic driver of terrestrial ecosystem processes. However, processes controlling interactions during and shortly after storms are often poorly seen and poorly sensed when direct observations are substituted with technological ones. We discuss how human observations complement technological ones and the benefits of scientists spending more time in the storm. Human observation can reveal ephemeral storm-related phenomena such as biogeochemical hot moments, organismal responses, and sedimentary processes that can then be explored in greater resolution using sensors and virtual experiments. Storm-related phenomena trigger lasting, oversized impacts on hydrologic and biogeochemical processes, organismal traits or functions, and ecosystem services at all scales. We provide examples of phenomena in forests, across disciplines and scales, that have been overlooked in past research to inspire mindful, holistic observation of ecosystems during storms. We conclude that technological observations alone are insufficient to trace the process complexity and unpredictability of fleeting biogeochemical or ecological events without the shower thoughts produced by scientists' human sensory and cognitive systems during storms.

Unlocking complex soil systems as carbon sinks: multi-pool management as the key.

Much research focuses on increasing carbon storage in mineral-associated organic matter (MAOM), in which carbon may persist for centuries to millennia. However, MAOM-targeted management is insufficient because the formation pathways of persistent soil organic matter are diverse and vary with environmental conditions. Effective management must also consider particulate organic matter (POM). In many soils, there is potential for enlarging POM pools, POM can persist over long time scales, and POM can be a direct precursor of MAOM. We present a framework for context-dependent management strategies that recognizes soils as complex systems in which environmental conditions constrain POM and MAOM formation.

Water availability dictates how plant traits predict demographic rates.

A major goal in ecology is to make generalizable predictions of organism responses to environmental variation based on their traits. However, straightforward relationships between traits and fitness are rare and likely to vary with environmental context. Characterizing how traits mediate demographic responses to the environment may enhance the predictions of organism responses to global change. We synthesized 15 years of demographic data and species-level traits in a shortgrass steppe to determine whether the effects of leaf and root traits on growth and survival depended on seasonal water availability. We predicted that (1) species with drought-tolerant traits, such as lower leaf turgor loss point (TLP) and higher leaf and root dry matter content (LDMC and RDMC), would be more likely to survive and grow in drier years due to higher wilting resistance, (2) these traits would not predict fitness in wetter years, and (3) traits that more directly measure physiological mechanisms of water use such as TLP would best predict demographic responses. We found that graminoids with more negative TLP and higher LDMC and RDMC had higher survival rates in drier years. Forbs demonstrated similar yet more variable responses. Graminoids grew larger in wetter years, regardless of traits. However, in both wet and dry years, graminoids with more negative TLP and higher LDMC and RDMC grew larger than less negative TLP and low LDMC and RDMC species. Traits significantly mediated the impact of drought on survival, but not growth, suggesting that survival could be a stronger driver of species' drought response in this system. TLP predicted survival in drier years, but easier to measure LDMC and RDMC were equal or better predictors. These results advance our understanding of the mechanisms by which drought drives population dynamics, and show that abiotic context determines how traits drive fitness.

Trading water for carbon in the future: Effects of elevated CO2 and warming on leaf hydraulic traits in a semiarid grassland.

The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO2 , warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre-dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO2 and/or warming. Effects of elevated CO2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO2 ; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO2 , which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near-term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture).

Contrasting responses of woody and grassland ecosystems to increased CO2 as water supply varies.

Experiments show that elevated atmospheric CO2 (eCO2) often enhances plant photosynthesis and productivity, yet this effect varies substantially and may be climate sensitive. Understanding if, where and how water supply regulates CO2 enhancement is critical for projecting terrestrial responses to increasing atmospheric CO2 and climate change. Here, using data from 14 long-term ecosystem-scale CO2 experiments, we show that the eCO2 enhancement of annual aboveground net primary productivity is sensitive to annual precipitation and that this sensitivity differs between woody and grassland ecosystems. During wetter years, CO2 enhancement increases in woody ecosystems but declines in grass-dominated systems. Consistent with this difference, woody ecosystems can increase leaf area index in wetter years more effectively under eCO2 than can grassland ecosystems. Overall, and across different precipitation regimes, woody systems had markedly stronger CO2 enhancement (24%) than grasslands (13%). We developed an empirical relationship to quantify aboveground net primary productivity enhancement on the basis of changes in leaf area index, providing a new approach for evaluating eCO2 impacts on the productivity of terrestrial ecosystems.

Plant traits related to precipitation sensitivity of species and communities in semiarid shortgrass prairie.

Understanding how plant communities respond to temporal patterns of precipitation in water-limited ecosystems is necessary to predict interannual variation and trends in ecosystem properties, including forage production, biogeochemical cycling, and biodiversity. In North American shortgrass prairie, we measured plant abundance, functional traits related to growth rate and drought tolerance, and aboveground net primary productivity to identify: species-level responsiveness to precipitation (precipitation sensitivity Sspp ) across functional groups; Sspp relationships to continuous plant traits; and whether continuous trait-Sspp relationships scaled to the community level. Across 32 plant species, we found strong bivariate relationships of both leaf dry matter content (LDMC) and leaf osmotic potential Ψosm with Sspp . Yet, LDMC and specific leaf area were retained in the lowest Akaike information criterion multiple regression model, explaining 59% of Sspp . Most relationships between continuous traits and Sspp scaled to the community level but were often contingent on the presence/absence of particular species and/or land management at a site. Thus, plant communities in shortgrass prairie may shift towards slower growing, more stress-resistant species in drought years and/or chronically drier climate. These findings highlight the importance of both leaf economic and drought tolerance traits in determining species and community responses to altered precipitation.

Extending the osmometer method for assessing drought tolerance in herbaceous species.

Community-scale surveys of plant drought tolerance are essential for understanding semi-arid ecosystems and community responses to climate change. Thus, there is a need for an accurate and rapid methodology for assessing drought tolerance strategies across plant functional types. The osmometer method for predicting leaf osmotic potential at full turgor (πo), a key metric of leaf-level drought tolerance, has resulted in a 50-fold increase in the measurement speed of this trait; however, the applicability of this method has only been tested in woody species and crops. Here, we assess the osmometer method for use in herbaceous grassland species and test whether πo is an appropriate plant trait for understanding drought strategies of herbaceous species as well as species distributions along climate gradients. Our model for predicting leaf turgor loss point (πTLP) from πo (πTLP = 0.80πo-0.845) is nearly identical to the model previously presented for woody species. Additionally, πo was highly correlated with πTLP for graminoid species (πtlp = 0.944πo-0.611; r2 = 0.96), a plant functional group previously flagged for having the potential to cause erroneous measurements when using an osmometer. We report that πo, measured with an osmometer, is well correlated with other traits linked to drought tolerance (namely, leaf dry matter content and leaf vulnerability to hydraulic failure) as well as climate extremes linked to water availability. The validation of the osmometer method in an herb-dominated ecosystem paves the way for rapid community-scale surveys of drought tolerance across plant functional groups, which could improve trait-based predictions of ecosystem responses to climate change.

Soil organic carbon stability in forests: Distinct effects of tree species identity and traits.

Rising atmospheric CO2 concentrations have increased interest in the potential for forest ecosystems and soils to act as carbon (C) sinks. While soil organic C contents often vary with tree species identity, little is known about if, and how, tree species influence the stability of C in soil. Using a 40 year old common garden experiment with replicated plots of eleven temperate tree species, we investigated relationships between soil organic matter (SOM) stability in mineral soils and 17 ecological factors (including tree tissue chemistry, magnitude of organic matter inputs to the soil and their turnover, microbial community descriptors, and soil physicochemical properties). We measured five SOM stability indices, including heterotrophic respiration, C in aggregate occluded particulate organic matter (POM) and mineral associated SOM, and bulk SOM δ15 N and ∆14 C. The stability of SOM varied substantially among tree species, and this variability was independent of the amount of organic C in soils. Thus, when considering forest soils as C sinks, the stability of C stocks must be considered in addition to their size. Further, our results suggest tree species regulate soil C stability via the composition of their tissues, especially roots. Stability of SOM appeared to be greater (as indicated by higher δ15 N and reduced respiration) beneath species with higher concentrations of nitrogen and lower amounts of acid insoluble compounds in their roots, while SOM stability appeared to be lower (as indicated by higher respiration and lower proportions of C in aggregate occluded POM) beneath species with higher tissue calcium contents. The proportion of C in mineral associated SOM and bulk soil ∆14 C, though, were negligibly dependent on tree species traits, likely reflecting an insensitivity of some SOM pools to decadal scale shifts in ecological factors. Strategies aiming to increase soil C stocks may thus focus on particulate C pools, which can more easily be manipulated and are most sensitive to climate change.

Elevated CO2 and water addition enhance nitrogen turnover in grassland plants with implications for temporal stability.

Temporal variation in soil nitrogen (N) availability affects growth of grassland communities that differ in their use and reuse of N. In a 7-year-long climate change experiment in a semi-arid grassland, the temporal stability of plant biomass production varied with plant N turnover (reliance on externally acquired N relative to internally recycled N). Species with high N turnover were less stable in time compared to species with low N turnover. In contrast, N turnover at the community level was positively associated with asynchrony in biomass production, which in turn increased community temporal stability. Elevated CO2 and summer irrigation, but not warming, enhanced community N turnover and stability, possibly because treatments promoted greater abundance of species with high N turnover. Our study highlights the importance of plant N turnover for determining the temporal stability of individual species and plant communities affected by climate change.

Grazing intensity differentially regulates ANPP response to precipitation in North American semiarid grasslands.

Grazing intensity elicits changes in the composition of plant functional groups in both shortgrass steppe (SGS) and northern mixed-grass prairie (NMP) in North America. How these grazing intensity-induced changes control aboveground net primary production (ANPP) responses to precipitation remains a central open question, especially in light of predicted climate changes. Here, we evaluated effects of four levels (none, light, moderate, and heavy) of long-term (>30 yr) grazing intensity in SGS and NMP on: (1) ANPP; (2) precipitation-use efficiency (PUE, ANPP : precipitation); and (3) precipitation marginal response (PMR; slope of a linear regression model between ANPP and precipitation). We advance prior work by examining: (1) the consequences of a range of grazing intensities (more grazed vs. ungrazed); and (2) how grazing-induced changes in ANPP and PUE are related both to shifts in functional group composition and physiological responses within each functional group. Spring (April-June) precipitation, the primary determinant of ANPP, was only 12% higher in NMP than in SGS, yet ANPP and PUE were 25% higher. Doubling grazing intensity in SGS and nearly doubling it in NMP reduced ANPP and PUE by only 24% and 33%, respectively. Increased grazing intensity reduced C3 graminoid biomass and increased C4 grass biomass in both grasslands. Functional group shifts affected PUE through biomass reductions, as PUE was positively associated with the relative abundance of C3 species and negatively with C4 species across both grasslands. At the community level, PMR was similar between grasslands and unaffected by grazing intensity. However, PMR of C3 graminoids in SGS was eightfold higher in the ungrazed treatment than under any grazed level. In NMP, PMR of C3 graminoids was only reduced under heavy grazing intensity. Knowing the ecological consequences of grazing intensity provides valuable information for mitigation and adaptation strategies in response to predicted climate change. For example, moderate grazing (the recommended rate) in SGS would sequester the same amount of aboveground carbon as light grazing because ANPP was nearly the same. In contrast, reductions in grazing intensity in NMP from moderate to light intensity would increase the amount of aboveground carbon sequestrated by 25% because of increased ANPP.

Integrating plant litter quality, soil organic matter stabilization, and the carbon saturation concept.

Labile, 'high-quality', plant litters are hypothesized to promote soil organic matter (SOM) stabilization in mineral soil fractions that are physicochemically protected from rapid mineralization. However, the effect of litter quality on SOM stabilization is inconsistent. High-quality litters, characterized by high N concentrations, low C/N ratios, and low phenol/lignin concentrations, are not consistently stabilized in SOM with greater efficiency than 'low-quality' litters characterized by low N concentrations, high C/N ratios, and high phenol/lignin concentrations. Here, we attempt to resolve these inconsistent results by developing a new conceptual model that links litter quality to the soil C saturation concept. Our model builds on the Microbial Efficiency-Matrix Stabilization framework (Cotrufo et al., 2013) by suggesting the effect of litter quality on SOM stabilization is modulated by the extent of soil C saturation such that high-quality litters are not always stabilized in SOM with greater efficiency than low-quality litters.

Effects of plant diversity, N fertilization, and elevated carbon dioxide on grassland soil N cycling in a long-term experiment.

The effects of global environmental changes on soil nitrogen (N) pools and fluxes have consequences for ecosystem functions such as plant productivity and N retention. In a 13-year grassland experiment, we evaluated how elevated atmospheric carbon dioxide (CO2 ), N fertilization, and plant species richness alter soil N cycling. We focused on soil inorganic N pools, including ammonium and nitrate, and two N fluxes, net N mineralization and net nitrification. In contrast with existing hypotheses, such as progressive N limitation, and with observations from other, often shorter, studies, elevated CO2 had relatively static and small, or insignificant, effects on soil inorganic N pools and fluxes. Nitrogen fertilization had inconsistent effects on soil N transformations, but increased soil nitrate and ammonium concentrations. Plant species richness had increasingly positive effects on soil N transformations over time, likely because in diverse subplots the concentrations of N in roots increased over time. Species richness also had increasingly positive effects on concentrations of ammonium in soil, perhaps because more carbon accumulated in soils of diverse subplots, providing exchange sites for ammonium. By contrast, subplots planted with 16 species had lower soil nitrate concentrations than less diverse subplots, especially when fertilized, probably due to greater N uptake capacity of subplots with 16 species. Monocultures of different plant functional types had distinct effects on N transformations and nitrate concentrations, such that not all monocultures differed from diverse subplots in the same manner. The first few years of data would not have adequately forecast the effects of N fertilization and diversity on soil N cycling in later years; therefore, the dearth of long-term manipulations of plant species richness and N inputs is a hindrance to forecasting the state of the soil N cycle and ecosystem functions in extant plant communities.

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.

Appraising the roles of nutrient availability, global change, and functional traits during the angiosperm rise to dominance.

To explain the rise of angiosperms during the Cretaceous, Berendse & Scheffer (Ecol. Lett., 12, 2009, 865) invoke feedbacks between leaf litter, soil nutrients, and growth, overlooking other factors affecting resource acquisition by Cretaceous plants. We evaluate their hypothesis, highlight alternative explanations, and emphasize use of a broader framework for understanding the angiosperm radiation.

Global patterns in leaf 13C discrimination and implications for studies of past and future climate.

Fractionation of carbon isotopes by plants during CO(2) uptake and fixation (Delta(leaf)) varies with environmental conditions, but quantitative patterns of Delta(leaf) across environmental gradients at the global scale are lacking. This impedes interpretation of variability in ancient terrestrial organic matter, which encodes climatic and ecological signals. To address this problem, we converted 3,310 published leaf delta(13)C values into mean Delta(leaf) values for 334 woody plant species at 105 locations (yielding 570 species-site combinations) representing a wide range of environmental conditions. Our analyses reveal a strong positive correlation between Delta(leaf) and mean annual precipitation (MAP; R(2) = 0.55), mirroring global trends in gross primary production and indicating stomatal constraints on leaf gas-exchange, mediated by water supply, are the dominant control of Delta(leaf) at large spatial scales. Independent of MAP, we show a lesser, negative effect of altitude on Delta(leaf) and minor effects of temperature and latitude. After accounting for these factors, mean Delta(leaf) of evergreen gymnosperms is lower (by 1-2.7 per thousand) than for other woody plant functional types (PFT), likely due to greater leaf-level water-use efficiency. Together, environmental and PFT effects contribute to differences in mean Delta(leaf) of up to 6 per thousand between biomes. Coupling geologic indicators of ancient precipitation and PFT (or biome) with modern Delta(leaf) patterns has potential to yield more robust reconstructions of atmospheric delta(13)C values, leading to better constraints on past greenhouse-gas perturbations. Accordingly, we estimate a 4.6 per thousand decline in the delta(13)C of atmospheric CO(2) at the onset of the Paleocene-Eocene Thermal Maximum, an abrupt global warming event approximately 55.8 Ma.

Accessibility of polybrominated diphenyl ether congeners in aging soil.

Polybrominated diphenyl ethers (PBDEs) are considered to be persistent environmental pollutants. Although soil is considered the most likely sink for these contaminants, little is known about the potential adsorption and aging of these compounds to the soil matrix. A previous study performed in our lab suggested that abiotic sorption of PBDEs to soil constituents was the most important determinant of PBDE accessibility. Building on this work, the present paper examined the availability of congeners of a commercial PBDE mixture (DE-71) in soils that varied in organic matter content, clay content, and pH. Both sterile and non-sterile soils were amended with DE-71 and then monitored over eight weeks of aging. Recovery of all congeners from soil by acetone extraction dropped significantly over time. Comparisons between sterile and non-sterile samples, along with the results of a separate phase partitioning study, indicated a strong affinity between soils and monitored DE-71 components. This general phenomenon was so dominant that, in this study, varying soil characteristics had no significant effect on PBDE recovery. Unexpectedly, final recovery was significantly higher in the non-sterile soil. The biological impact on congener accessibility was also observed when zucchini and radish plants were grown in soil that had been aged 8 weeks following DE-71 fortification. After 10 weeks of growth, recovery of congeners was up to five times higher than it had been prior to planting.

Polybrominated diphenyl ethers: causes for concern and knowledge gaps regarding environmental distribution, fate and toxicity.

This manuscript critically considers several areas of study of the polybrominated diphenyl ether compounds. Specifically, a brief review of PBDE toxicity is followed by an in depth discussion of PBDE occurrence in abiotic and biotic environmental matrices. Temporal and geographic trends are examined in conjunction with risk assessment factors. Rather than summarize or tabulate the growing body of literature on PBDEs in the environment, the overall goal of this review paper is to highlight broad patterns that may contribute to a more holistic understanding of PBDE behavior in the environment, as well as to identify critical areas of research that warrant further attention.