Comparative venation costs of monocotyledon and dicotyledon species in the eastern Colorado steppe.

MAIN CONCLUSION: Leaf vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles did not differ between monocot and dicot species in 21 species from the eastern Colorado steppe. Dicots possessed significantly larger minor vein networks than monocots. Across the tree of life, there is evidence that dendritic vascular transport networks are optimized, balancing maximum speed and integrity of resource delivery with minimal resource investment in transport and infrastructure. Monocot venation, however, is not dendritic, and remains parallel down to the smallest vein orders with no space-filling capillary networks. Given this departure from the "optimized" dendritic network, one would assume that monocots are operating at a significant energetic disadvantage. In this study, we investigate whether monocot venation networks bear significantly greater carbon/construction costs per leaf volume than co-occurring dicots in the same ecosystem, and if so, what physiological or ecological advantage the monocot life form possesses to compensate for this deficit. Given that venation networks could also be optimized for leaf mechanical support or provide herbivory defense, we measured the vascular system of both monocot and dicots at three scales to distinguish between leaf investment in mechanical support (macroscopic vein), total transport and capacitance (vascular bundle), or exclusively water transport (xylem) for both parallel and dendritic venation networks. We observed that vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles was not significantly different between monocot species and dicot species. Dicots, however, possess significantly larger minor vein networks than monocots. The 19 species subjected to gas-exchange measurement in the field displayed a broad range of Amax and but demonstrated no significant relationships with any metric of vascular network size in major or minor vein classes. Given that monocots do not seem to display any leaf hydraulic disadvantage relative to dicots, it remains an important research question why parallel venation (truly parallel, down to the smallest vessels) has not arisen more than once in the history of plant evolution.

Grass veins are leaky pipes: vessel widening in grass leaves explain variation in stomatal conductance and vessel diameter among species.

The widening of xylem vessels from tip to base of trees is an adaptation to minimize the hydraulic resistance of a long pathway. Given that parallel veins of monocot leaves do not branch hierarchically, vessels should also widen basipetally but, in addition to minimizing resistance, should also account for water volume lost to transpiration since they supply water to the lamina along their lengths, that is 'leakiness'. We measured photosynthesis, stomatal conductance, and vessel diameter at five locations along each leaf of five perennial grass species. We found that the rate of conduit widening in grass leaves was larger than the widening exponent required to minimize pathlength resistance (0.35 vs c. 0.22). Furthermore, variation in the widening exponent among species was positively correlated with maximal stomatal conductance (r2 = 0.20) and net CO2 assimilation (r2 = 0.45). These results suggest that faster rates of conduit widening (> 0.22) were associated with higher rates of water loss. Taken together, our results show that the widening exponent is linked to plant function in grass leaves and that natural selection has favored parallel vein networks that are constructed to meet transpiration requirements while minimizing hydraulic resistance within grass blades.

Physiological trait networks enhance understanding of crop growth and water use in contrasting environments.

Plant function arises from a complex network of structural and physiological traits. Explicit representation of these traits, as well as their connections with other biophysical processes, is required to advance our understanding of plant-soil-climate interactions. We used the Terrestrial Regional Ecosystem Exchange Simulator (TREES) to evaluate physiological trait networks in maize. Net primary productivity (NPP) and grain yield were simulated across five contrasting climate scenarios. Simulations achieving high NPP and grain yield in high precipitation environments featured trait networks conferring high water use strategies: deep roots, high stomatal conductance at low water potential ("risky" stomatal regulation), high xylem hydraulic conductivity and high maximal leaf area index. In contrast, high NPP and grain yield was achieved in dry environments with low late-season precipitation via water conserving trait networks: deep roots, high embolism resistance and low stomatal conductance at low leaf water potential ("conservative" stomatal regulation). We suggest that our approach, which allows for the simultaneous evaluation of physiological traits, soil characteristics and their interactions (i.e., networks), has potential to improve our understanding of crop performance in different environments. In contrast, evaluating single traits in isolation of other coordinated traits does not appear to be an effective strategy for predicting plant performance.

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).

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.

Genetic and functional variation across regional and local scales is associated with climate in a foundational prairie grass.

Global change forecasts in ecosystems require knowledge of within-species diversity, particularly of dominant species within communities. We assessed site-level diversity and capacity for adaptation in Bouteloua gracilis, the dominant species in the Central US shortgrass steppe biome. We quantified genetic diversity from 17 sites across regional scales, north to south from New Mexico to South Dakota, and local scales in northern Colorado. We also quantified phenotype and plasticity within and among sites and determined the extent to which phenotypic diversity in B. gracilis was correlated with climate. Genome sequencing indicated pronounced population structure at the regional scale, and local differences indicated that gene flow and/or dispersal may also be limited. Within a common environment, we found evidence of genetic divergence in biomass-related phenotypes, plasticity, and phenotypic variance, indicating functional divergence and different adaptive potential. Phenotypes were differentiated according to climate, chiefly median Palmer Hydrological Drought Index and other aridity metrics. Our results indicate conclusive differences in genetic variation, phenotype, and plasticity in this species and suggest a mechanism explaining variation in shortgrass steppe community responses to global change. This analysis of B. gracilis intraspecific diversity across spatial scales will improve conservation and management of the shortgrass steppe ecosystem in the future.

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.

Vessel scaling in evergreen angiosperm leaves conforms with Murray's law and area-filling assumptions: implications for plant size, leaf size and cold tolerance.

Water transport in leaf vasculature is a fundamental process affecting plant growth, ecological interactions and ecosystem productivity, yet the architecture of leaf vascular networks is poorly understood. Although Murray's law and the West-Brown-Enquist (WBE) theories predict convergent scaling of conduit width and number, it is not known how conduit scaling is affected by habitat aridity or temperature. We measured the scaling of leaf size, conduit width and conduit number within the leaves of 36 evergreen Angiosperms spanning a large range in aridity and temperature in eastern Australia. Scaling of conduit width and number in midribs and 2° veins did not differ across species and habitats (P > 0.786), and did not differ from that predicted by Murray's law (P = 0.151). Leaf size was strongly correlated with the hydraulic radius of petiole conduits (r2  = 0.83, P < 0.001) and did not differ among habitats (P > 0.064), nor did the scaling exponent differ significantly from that predicted by hydraulic theory (P = 0.086). The maximum radius of conduits in petioles was positively correlated with the temperature of the coldest quarter (r2  = 0.67; P < 0.001), suggesting that habitat temperature restricts the occurrence of wide-conduit species in cold habitats.

A safety vs efficiency trade-off identified in the hydraulic pathway of grass leaves is decoupled from photosynthesis, stomatal conductance and precipitation.

A common theme in plant physiological research is the trade-off between stress tolerance and growth; an example of this trade-off at the tissue level is the safety vs efficiency hypothesis, which suggests that plants with the greatest resistance to hydraulic failure should have low maximum hydraulic conductance. Here, we quantified the leaf-level drought tolerance of nine C4 grasses as the leaf water potential at which plants lost 50% (P50 × RR ) of maximum leaf hydraulic conductance (Ksat ), and compared this trait with other leaf-level and whole-plant functions. We found a clear trade-off between Ksat and P50 × RR when Ksat was normalized by leaf area and mass (P = 0.05 and 0.01, respectively). However, no trade-off existed between P50 × RR and gas-exchange rates; rather, there was a positive relationship between P50 × RR and photosynthesis (P = 0.08). P50 × RR was not correlated with species distributions based on precipitation (P = 0.70), but was correlated with temperature during the wettest quarter of the year (P < 0.01). These results suggest a trade-off between safety and efficiency in the hydraulic system of grass leaves, which can be decoupled from other leaf-level functions. The unique physiology of C4 plants and adaptations to pulse-driven systems may provide mechanisms that could decouple hydraulic conductance from other plant functions.

Beyond arctic and alpine: the influence of winter climate on temperate ecosystems.

Winter climate is expected to change under future climate scenarios, yet the majority of winter ecology research is focused in cold-climate ecosystems. In many temperate systems, it is unclear how winter climate relates to biotic responses during the growing season. The objective of this study was to examine how winter weather relates to plant and animal communities in a variety of terrestrial ecosystems ranging from warm deserts to alpine tundra. Specifically, we examined the association between winter weather and plant phenology, plant species richness, consumer abundance, and consumer richness in 11 terrestrial ecosystems associated with the U.S. Long-Term Ecological Research (LTER) Network. To varying degrees, winter precipitation and temperature were correlated with all biotic response variables. Bud break was tightly aligned with end of winter temperatures. For half the sites, winter weather was a better predictor of plant species richness than growing season weather. Warmer winters were correlated with lower consumer abundances in both temperate and alpine systems. Our findings suggest winter weather may have a strong influence on biotic activity during the growing season and should be considered in future studies investigating the effects of climate change on both alpine and temperate systems.

Lack of eutrophication in a tallgrass prairie ecosystem over 27 years.

Many North American grasslands are receiving atmospheric nitrogen (N) deposition at rates above what are considered critical eutrophication thresholds. Yet, potential changes in grassland function due to anthropogenic N deposition are poorly resolved, especially considering that other dynamic factors such as land use and precipitation can also affect N availability. To better understand whether elevated N deposition has altered ecosystem structure or function in North American grasslands, we analyzed a 27-year record of ecophysiological, community, and ecosystem metrics for an annually burned Kansas tallgrass prairie. Over this time, despite increasing rates of N deposition that are within the range of critical loads for grasslands, there was no evidence of eutrophication. Plant N concentrations did not increase, soil moisture did not decline, forb diversity did not decline, and the relative abundance of dominant grasses did not shift toward more eutrophic species. Neither aboveground primary productivity nor N availability to plants increased. The fates of deposited N in grasslands are still uncertain, and could include management losses through burning and grazing. However, evidence from this grassland indicates that eutrophication of North American grassland ecosystems is not an inevitable consequence of current levels of N deposition.

Elevated CO2 counteracts effects of water stress on woody rangeland-encroaching species.

The ubiquity of woody plant expansion across many rangelands globally has led to the hypothesis that the global rise in atmospheric carbon dioxide concentration ([CO2]) is a global driver facilitating C3 woody plant expansion. Increasing [CO2] also influences precipitation patterns seasonally and across the landscape, which often results in the prevalence of drought in rangelands. To test the potential for [CO2] to facilitate woody plant growth, we conducted a greenhouse study for 150 days to measure CO2 effects on juveniles from four woody species (C. drummondii, R. glabra, G. triacanthos and J. osteosperma) that are actively expanding into rangelands of North America. We assessed chronic water-stress (nested within CO2 treatments) and its interaction with elevated [CO2] (800 ppm) on plant growth physiology for 84 days. We measured leaf-level gas exchange, tissue-specific starch concentrations and biomass. We found that elevated [CO2] increased photosynthetic rates, intrinsic water-use efficiencies, and leaf starch concentrations in all woody species but at different rates and concentrations. Elevated [CO2] increased leaf starch levels for C. drummondii, G. triacanthos, J. osteosperma, and R. glabra by 90%, 39%, 68%, and 41% respectively. We also observed that elevated [CO2] ameliorated the physiological effects of chronic water stress for all our juvenile woody species within this study. Elevated [CO2] diminished the impact of water stress on the juvenile plants, potentially alleviating an abiotic limitation to woody plant establishment in rangelands, thus facilitating the expansion of woody plants in the future.

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.

Natural selection drives clinal life history patterns in the perennial sunflower species, Helianthus maximiliani.

In plants, ecologically important life history traits often display clinal patterns of population divergence. Such patterns can provide strong evidence for spatially varying selection across environmental gradients but also may result from nonselective processes, such as genetic drift, population bottlenecks and spatially restricted gene flow. Comparison of population differentiation in quantitative traits (measured as Q(ST) ) with neutral molecular markers (measured as F(ST) ) provides a useful tool for understanding the relative importance of adaptive and nonadaptive processes in the formation and maintenance of clinal variation. Here, we demonstrate the existence of geographic variation in key life history traits in the diploid perennial sunflower species Helianthus maximiliani across a broad latitudinal transect in North America. Strong population differentiation was found for days to flowering, growth rate and multiple size-related traits. Differentiation in these traits greatly exceeds neutral predictions, as determined both by partial Mantel tests and by comparisons of global Q(ST) values with theoretical F(ST) distributions. These findings indicate that clinal variation in these life history traits likely results from local adaptation driven by spatially heterogeneous environments.

Linking plant growth responses across topographic gradients in tallgrass prairie.

Aboveground biomass in grasslands varies according to landscape gradients in resource availability and seasonal patterns of growth. Using a transect spanning a topographic gradient in annually burned ungrazed tallgrass prairie, we measured changes in the height of four abundant C(4) grass species, LAI, biomass, and cumulative carbon flux using two closely located eddy flux towers. We hypothesized that seasonal patterns of plant growth would be similar across the gradient, but the magnitude of growth and biomass accumulation would vary by topographic position, reflecting spatial differences in microclimate, slope, elevation, and soil depth. Thus, identifying and measuring local growth responses according to topographic variability should significantly improve landscape predictions of aboveground biomass. For most of the growth variables measured, classifying topography into four positions best captured the inherent spatial variability. Biomass produced, seasonal LAI and species height increased from the upland and break positions to the slope and lowland. Similarly, cumulative carbon flux in 2008 was greater in lowland versus upland tower locations (difference of 64 g m(-2) by DOY 272). Differences in growth by topographic position reflected increased production of flowering culms by Andropogon gerardii and Sorghastrum nutans in lowland. Varying growth responses by these species may be a significant driver of biomass and carbon flux differences by topographic position, at least for wet years. Using a digital elevation model to classify the watershed into topographic positions, we performed a geographically weighted regression to predict landscape biomass. The minimum and maximum predictions of aboveground biomass for this watershed had a large range (86-393 t per 40.4 ha), illustrating the drastic spatial variability in growth within this annually-burned grassland.

Local adaptation to precipitation in the perennial grass Elymus elymoides: Trade-offs between growth and drought resistance traits.

Understanding local adaptation to climate is critical for managing ecosystems in the face of climate change. While there have been many provenance studies in trees, less is known about local adaptation in herbaceous species, including the perennial grasses that dominate arid and semiarid rangeland ecosystems. We used a common garden study to quantify variation in growth and drought resistance traits in 99 populations of Elymus elymoides from a broad geographic and climatic range in the western United States. Ecotypes from drier sites produced less biomass and smaller seeds, and had traits associated with greater drought resistance: small leaves with low osmotic potential and high integrated water use efficiency (δ13C). Seasonality also influenced plant traits. Plants from regions with relatively warm, wet summers had large seeds, large leaves, and low δ13C. Irrespective of climate, we also observed trade-offs between biomass production and drought resistance traits. Together, these results suggest that much of the phenotypic variation among E. elymoides ecotypes represents local adaptation to differences in the amount and timing of water availability. In addition, ecotypes that grow rapidly may be less able to persist under dry conditions. Land managers may be able to use this variation to improve restoration success by seeding ecotypes with multiple drought resistance traits in areas with lower precipitation. The future success of this common rangeland species will likely depend on the use of tools such as seed transfer zones to match local variation in growth and drought resistance to predicted climatic conditions.

Thirteen decades of foliar isotopes indicate declining nitrogen availability in central North American grasslands.

*Humans are increasing both the deposition of reactive nitrogen (N) and concentrations of atmospheric CO(2) on Earth, but the combined effects on terrestrial ecosystems are not clear. In the absence of historical records, it is difficult to know if N availability is currently increasing or decreasing on regional scales. *To determine the nature and timing of past changes in grassland ecosystem dynamics, we measured the composition of stable carbon (C) and N isotopes in leaf tissue from 545 herbarium specimens of 24 vascular plant species collected in Kansas, USA from 1876 to 2008. We also parameterized a simple model of the terrestrial N cycle coupled with a stable isotope simulator to constrain processes consistent with observed patterns. *A prolonged decline in foliar N concentrations began in 1926, while a prolonged decline in foliar delta(15)N values began in 1940. Changes in the difference between foliar and atmospheric C isotopes reveal slightly increased photosynthetic water use efficiency since 1876. *The declines in foliar N concentrations and foliar delta(15)N suggest declining N availability in these grasslands during the 20th century despite decades of anthropogenic N deposition. Our results are consistent with progressive-nitrogen-limitation-type hypotheses where declines in N availability are driven by increased ecosystem N storage as a result of increased atmospheric CO(2).

Using nocturnal cold air drainage flow to monitor ecosystem processes in complex terrain.

This paper presents initial investigations of a new approach to monitor ecosystem processes in complex terrain on large scales. Metabolic processes in mountainous ecosystems are poorly represented in current ecosystem monitoring campaigns because the methods used for monitoring metabolism at the ecosystem scale (e.g., eddy covariance) require flat study sites. Our goal was to investigate the potential for using nocturnal down-valley winds (cold air drainage) for monitoring ecosystem processes in mountainous terrain from two perspectives: measurements of the isotopic composition of ecosystem-respired CO2 (delta13C(ER)) and estimates of fluxes of CO2 transported in the drainage flow. To test if this approach is plausible, we monitored the wind patterns, CO2 concentrations, and the carbon isotopic composition of the air as it exited the base of a young (approximately 40 yr-old) and an old (>450 yr-old) steeply sided Douglas-fir watershed. Nocturnal cold air drainage within these watersheds was strong, deep, and occurred on more than 80% of summer nights. The depth of cold air drainage rapidly increased to tower height or greater when the net radiation at the top of the tower approached zero. The carbon isotope composition of CO2 in the drainage system holds promise as an indicator of variation in basin-scale physiological processes. Although there was little vertical variation in CO2 concentration at any point in time, we found that the range of CO2 concentration over a single evening was sufficient to estimate delta 13C(ER) from Keeling plot analyses. The seasonal variation in delta 13C(ER) followed expected trends: during the summer dry season delta 13C(ER) became less negative (more enriched in 13C), but once rain returned in the fall, delta 13C(ER) decreased. However, we found no correlation between recent weather (e.g., vapor pressure deficit) and delta 13C(ER) either concurrently or with up to a one-week lag. Preliminary estimates suggest that the nocturnal CO2 flux advecting past the 28-m tower is a rather small fraction (<20%) of the watershed-scale respiration. This study demonstrates that monitoring the isotopic composition and CO2 concentration of cold air drainage at the base of a watershed provides a new tool for quantifying ecosystem metabolism in mountainous ecosystems on the basin scale.

Partitioning hydraulic resistance in Sorghum bicolor leaves reveals unique correlations with stomatal conductance during drought.

The hydraulic architecture of leaves represents the final path along which liquid water travels through the plant and comprises a significant resistance for water movement, especially for grasses. We partitioned leaf hydraulic resistance of six genotypes of Sorghum bicolor L. (Moench) into leaf specific hydraulic resistance within the large longitudinal veins (r*LV) and outside the large veins (r*OLV), and correlated these resistances with the response of stomatal conductance (gs) and photosynthesis (A) to drought. Under well-watered conditions, gs was tightly correlated with r*OLV (r2=0.95), but as soil moisture decreased, gs was more closely correlated with r*LV (r2=0.97). These results suggest that r*OLV limits maximum rates of gas exchange, but the ability to efficiently move water long distances (low r*LV) becomes more important for the maintenance of cell turgor and gas exchange as soil moisture declines. Hydraulic resistance through the leaf was negatively correlated with evapotranspiration (P<0.001) resulting in more conservative water use in genotypes with large leaf resistance. These results illustrate the functional significance of leaf resistance partitioning to declining soil moisture in a broadly-adapted cereal species.

Evidence of physiological decoupling from grassland ecosystem drivers by an encroaching woody shrub.

Shrub encroachment of grasslands is a transformative ecological process by which native woody species increase in cover and frequency and replace the herbaceous community. Mechanisms of encroachment are typically assessed using temporal data or experimental manipulations, with few large spatial assessments of shrub physiology. In a mesic grassland in North America, we measured inter- and intra-annual variability in leaf δ(13)C in Cornus drummondii across a grassland landscape with varying fire frequency, presence of large grazers and topographic variability. This assessment of changes in individual shrub physiology is the largest spatial and temporal assessment recorded to date. Despite a doubling of annual rainfall (in 2008 versus 2011), leaf δ(13)C was statistically similar among and within years from 2008-11 (range of -28 to -27‰). A topography*grazing interaction was present, with higher leaf δ(13)C in locations that typically have more bare soil and higher sensible heat in the growing season (upland topographic positions and grazed grasslands). Leaf δ(13)C from slopes varied among grazing contrasts, with upland and slope leaf δ(13)C more similar in ungrazed locations, while slopes and lowlands were more similar in grazed locations. In 2011, canopy greenness (normalized difference vegetation index - NDVI) was assessed at the centroid of individual shrubs using high-resolution hyperspectral imagery. Canopy greenness was highest mid-summer, likely reflecting temporal periods when C assimilation rates were highest. Similar to patterns seen in leaf δ(13)C, NDVI was highest in locations that typically experience lowest sensible heat (lowlands and ungrazed). The ability of Cornus drummondii to decouple leaf physiological responses from climate variability and fire frequency is a likely contributor to the increase in cover and frequency of this shrub species in mesic grassland and may be generalizable to other grasslands undergoing woody encroachment.