Rapid increases in shrubland and forest intrinsic water-use efficiency during an ongoing megadrought.

Globally, intrinsic water-use efficiency (iWUE) has risen dramatically over the past century in concert with increases in atmospheric CO2 concentration. This increase could be further accelerated by long-term drought events, such as the ongoing multidecadal "megadrought" in the American Southwest. However, direct measurements of iWUE in this region are rare and largely constrained to trees, which may bias estimates of iWUE trends toward more mesic, high elevation areas and neglect the responses of other key plant functional types such as shrubs that are dominant across much of the region. Here, we found evidence that iWUE is increasing in the Southwest at one of the fastest rates documented due to the recent drying trend. These increases were particularly large across three common shrub species, which had a greater iWUE sensitivity to aridity than Pinus ponderosa, a common tree species in the western United States. The sensitivity of both shrub and tree iWUE to variability in atmospheric aridity exceeded their sensitivity to increasing atmospheric [CO2]. The shift to more water-efficient vegetation would be, all else being equal, a net positive for plant health. However, ongoing trends toward lower plant density, diminished growth, and increasing vegetation mortality across the Southwest indicate that this increase in iWUE is unlikely to offset the negative impacts of aridification.

Examination of amino acid hydrogen isotope measurements of scalp hair for region-of-origin studies.

RATIONALE: Hydrogen isotope (δ2 H) analysis of keratinaceous bulk tissues has been used in forensic science to reconstruct an individual's travel history or determine their region-of-origin. Here, we use a compound-specific approach to examine patterns of individual amino acid δ2 H values in relation to those of local tap water, bulk scalp hair tissues, and region-of-origin. METHODS: We measured δ2 H values of amino acids in anonymously collected scalp hair (n = 67) and tap water from 28 locations in the United States. Samples were hydrolyzed into their constituent amino acids, derivatized alongside in-house reference materials, and analyzed in triplicate using a GC-C-IRMS system. RESULTS: Non-essential amino acid (AANESS ) δ2 H values and their corresponding tap water samples varied systematically across continental regions. Hydrogen isotope values of alanine, glutamic acid, and glycine were significantly correlated with tap water and an estimated 42%-51% of the hydrogen atoms in these AANESS were derived from tap water. We used linear discriminate analysis (LDA) to explore regional patterns in scalp hair bulk tissue and amino acid δ2 H values. For the model that included AANESS data, 87% of the variance was explained by the first linear discriminant axis (LD1), and was driven by bulk hair tissue, alanine, and proline. This model had an overall 72% successful reclassification with samples from the south and northwest regions reclassifying correctly 92% and 78% of the time, respectively. For the model that included AAESS data, LD1 explained 81% of the variation and was driven bulk hair, threonine, valine, phenylalanine, and isoleucine. The overall reclassification rate for the model that included AAESS was 70%. CONCLUSIONS: Our findings suggest that δ2 H analyses of AANESS and AAESS could help improve geolocation models for human and wildlife forensics by simultaneously providing information about both dietary and tap water inputs of hydrogen to tissue synthesis.

The carbon balance of plants: economics, optimization, and trait spectra in a historical perspective.

Over fifty years have passed since the publication of Harold Mooney's formative paper, "The Carbon Balance of Plants" on pages 315-346 of Volume 3 (1972) of Annual Review of Ecology and Systematics. Arguably, the conceptual framework presented in that paper, and the work by Mooney and his students leading up to the paper, provided the foundational principles from which core disciplines emerged in plant economic theory, functional trait theory and, more generally, plant physiological ecology. Here, we revisit the primary impacts of those early discoveries to understand how researchers constructed major concepts in our understanding of plant adaptations, and where those concepts are likely to take us in the near future. The discipline of functional trait ecology, which is rooted in the principles of evolutionary and economic optimization, has captured the imagination of the plant physiological ecology research community, though its emphasis has shifted toward predicting species distributions and ecological roles across resource gradients. In the face of 'big-data' research pursuits that are revealing trait expression patterns at the cellular level and mass and energy exchange patterns at the planetary scale, an opportunity exists to reconnect the principles of plant carbon balance and evolutionary optimization with trait origins at the genetic and cellular scales and trait impacts at the global scale.

Multidecadal records of intrinsic water-use efficiency in the desert shrub Encelia farinosa reveal strong responses to climate change.

While tree rings have enabled interannual examination of the influence of climate on trees, this is not possible for most shrubs. Here, we leverage a multidecadal record of annual foliar carbon isotope ratio collections coupled with 39 y of survey data from two populations of the drought-deciduous desert shrub Encelia farinosa to provide insight into water-use dynamics and climate. This carbon isotope record provides a unique opportunity to examine the response of desert shrubs to increasing temperature and water stress in a region where climate is changing rapidly. Population mean carbon isotope ratios fluctuated predictably in response to interannual variations in temperature, vapor pressure deficit, and precipitation, and responses were similar among individuals. We leveraged the well-established relationships between leaf carbon isotope ratios and the ratio of intracellular to ambient CO2 concentrations to calculate intrinsic water-use efficiency (iWUE) of the plants and to quantify plant responses to long-term environmental change. The population mean iWUE value increased by 53 to 58% over the study period, much more than the 20 to 30% increase that has been measured in forests [J. Peñuelas, J. G. Canadell, R. Ogaya, Glob. Ecol. Biogeogr. 20, 597-608 (2011)]. Changes were associated with both increased CO2 concentration and increased water stress. Individuals whose lifetimes spanned the entire study period exhibited increases in iWUE that were very similar to the population mean, suggesting that there was significant plasticity within individuals rather than selection at the population scale.

Stable isotopes in hair reveal dietary protein sources with links to socioeconomic status and health.

Carbon and nitrogen isotope ratios in hair sampled from 65 communities across the central and intermountain regions of the United States and more intensively throughout 29 ZIP codes in the Salt Lake Valley, Utah, revealed a dietary divergence related to socioeconomic status as measured by cost of living, household income, and adjusted gross income. Corn-fed, animal-derived proteins were more common in the diets of lower socioeconomic status populations than were plant-derived proteins, with individual estimates of animal-derived protein diets as high as 75%; United States towns and cities averaged 57%. Similar patterns were seen across the socioeconomic status spectrum in the Salt Lake Valley. It is likely that corn-fed animal proteins were associated with concentrated animal-feeding operations, a common practice for industrial animal production in the United States today. Given recent studies highlighting the negative impacts of animal-derived proteins in our diets, hair carbon isotope ratios could provide an approach for scaling assessments of animal-sourced foods and health risks in communities across the United States.

Intrinsic water-use efficiency influences establishment in Encelia farinosa.

We describe establishment of Encelia farinosa, a drought-deciduous shrub common to the Mojave and Sonoran Deserts, based on annual observations of two populations between 1980 and 2020. Only 11 establishment events of 50 + yearlings (0.02-0.03 individuals m-2) occurred during this monitoring period; in 68% of the years fewer than 10 yearlings were established. Yearling survival to adulthood (age 4) ranged from 88 to 5% and was significantly related to cumulative precipitation. Juvenile survival rates were lowest during the current megadrought period. We calculated intrinsic water-use efficiency (iWUE) and observed the widest variations in iWUE values among the youngest plants. Among juveniles, surviving yearlings with the lowest iWUE values exhibited upward ontogenetic shifts in iWUE values, whereas those yearlings with the highest initial iWUE values exhibited little if any change. Juvenile size, higher iWUE values, and greater likelihood of surviving were all positively related with each other over the past several decades. Furthermore, iWUE and photosynthetic capacity were positively related to each other, providing a mechanistic explanation for why increased iWUE values among juveniles could lead to greater survival rates and to larger plants under water-deficit conditions. We posit that there is bi-directional selection for genotypic variations in iWUE values among E. farinosa and that this variation is selected for because of interannual environmental heterogeneity in precipitation and VPD associated with both high- and low-frequency climate cycles. Extreme drought cycles may favor plants with higher iWUE values, whereas more mesic periods may allow for greater persistence of lower iWUE genotypes.

Heterogeneous isotope effects decouple conifer leaf and branch sugar δ18O and δ13C.

Isotope ratios of tree-ring cellulose are a prominent tool to reconstruct paleoclimate and plant responses to environmental variation. Current models for cellulose isotope ratios assume a transfer of the environmental signals recorded in bulk leaf water to carbohydrates and ultimately into stem cellulose. However, the isotopic signal of carbohydrates exported from leaf to branch may deviate from mean leaf values if spatial heterogeneity in isotope ratios exists in the leaf. We tested whether the isotopic heterogeneity previously observed along the length of a ponderosa pine (Pinus ponderosa) leaf water was preserved in photosynthetic products. We observed an increase in both sugar and bulk tissue δ18O values along the needle, but the increase in carbohydrate δ18O values was dampened relative to the trend observed in leaf water. In contrast, δ13C values of both sugar and bulk organic matter were invariant along the needle. Phloem-exported sugar measured in the branch below the needles did not match whole-needle values of δ18O or δ13C. Instead, there was a near-constant offset observed between the branch and needle sugar δ13C values, while branch δ18O values were most similar to δ18O values observed for sugar at the base of the needle. The observed offset between the branch and needle sugar δ18O values likely arises from partial isotope oxygen exchange between sugars and water during phloem loading and transport. An improved understanding of the conditions producing differential δ13C and δ18O isotope effects between branch phloem and needle sugars could improve tree-ring-based climate reconstructions.

Long-term nitrogen isotope dynamics in Encelia farinosa reflect plant demographics and climate.

While plant δ15 N values have been applied to understand nitrogen (N) dynamics, uncertainties regarding intraspecific and temporal variability currently limit their application. We used a 28 yr record of δ15 N values from two Mojave Desert populations of Encelia farinosa to clarify sources of population-level variability. We leveraged > 3500 foliar δ15 N observations collected alongside structural, physiological, and climatic data to identify plant and environmental contributors to δ15 N values. Additional sampling of soils, roots, stems, and leaves enabled assessment of the distribution of soil N content and δ15 N, intra-plant fractionations, and relationships between soil and plant δ15 N values. We observed extensive within-population variability in foliar δ15 N values and found plant age and foliar %N to be the strongest predictors of individual δ15 N values. There were consistent differences between root, stem, and leaf δ15 N values (spanning c. 3‰), but plant and bulk soil δ15 N values were unrelated. Plant-level variables played a strong role in influencing foliar δ15 N values, and interannual relationships between climate and δ15 N values were counter to previously recognized spatial patterns. This long-term record provides insights regarding the interpretation of δ15 N values that were not available from previous large-scale syntheses, broadly enabling more effective application of foliar δ15 N values.

Seasonal and diurnal trends in progressive isotope enrichment along needles in two pine species.

The Craig-Gordon type (C-G) leaf water isotope enrichment models assume a homogeneous distribution of enriched water across the leaf surface, despite observations that Δ18 O can become increasingly enriched from leaf base to tip. Datasets of this 'progressive isotope enrichment' are limited, precluding a comprehensive understanding of (a) the magnitude and variability of progressive isotope enrichment, and (b) how progressive enrichment impacts the accuracy of C-G leaf water model predictions. Here, we present observations of progressive enrichment in two conifer species that capture seasonal and diurnal variability in environmental conditions. We further examine which leaf water isotope models best capture the influence of progressive enrichment on bulk needle water Δ18 O. Observed progressive enrichment was large and equal in magnitude across both species. The magnitude of this effect fluctuated seasonally in concert with vapour pressure deficit, but was static in the face of diurnal cycles in meteorological conditions. Despite large progressive enrichment, three variants of the C-G model reasonably successfully predicted bulk needle Δ18 O. Our results thus suggest that the presence of progressive enrichment does not impact the predictive success of C-G models, and instead yields new insight regarding the physiological and anatomical mechanisms that cause progressive isotope enrichment.

Interactions among intrinsic water-use efficiency and climate influence growth and flowering in a common desert shrub.

Plants make leaf-level trade-offs between photosynthetic carbon assimilation and water loss, and the optimal balance between the two is dependent, in part, on water availability. "Conservative" water-use strategies, in which minimizing water loss is prioritized over assimilating carbon, tend to be favored in arid environments, while "aggressive" water-use strategies, in which carbon assimilation is prioritized over water conservation, are often favored in mesic environments. When derived from foliar carbon isotope ratios, intrinsic water-use efficiency (iWUE) serves as a seasonally integrated indicator of the balance of carbon assimilation to water loss at the leaf level. Here, we used a multi-decadal record of annual iWUE, growth, and flowering from a single population of Encelia farinosa in the Mojave Desert to evaluate the effect of iWUE on plant performance across interannual fluctuations in water availability. We identified substantial variability in iWUE among individuals and found that iWUE interacted with water availability to significantly influence growth and flowering. However, the relationships between iWUE, water availability, and plant performance did not universally suggest that "conservative" water-use strategies were advantageous in dry years or that "aggressive" strategies were advantageous in wet years. iWUE was positively related to the odds of growth regardless of water availability and to the odds of flowering in dry years, but negatively related to growth rates in dry years. In addition, we found that leaf nitrogen content affected interannual plant performance and that an individual's iWUE plasticity in response to fluctuations in aridity was negatively related to early life drought survival and growth.

Long-term urban carbon dioxide observations reveal spatial and temporal dynamics related to urban characteristics and growth.

Cities are concentrated areas of CO2 emissions and have become the foci of policies for mitigation actions. However, atmospheric measurement networks suitable for evaluating urban emissions over time are scarce. Here we present a unique long-term (decadal) record of CO2 mole fractions from five sites across Utah's metropolitan Salt Lake Valley. We examine "excess" CO2 above background conditions resulting from local emissions and meteorological conditions. We ascribe CO2 trends to changes in emissions, since we did not find long-term trends in atmospheric mixing proxies. Three contrasting CO2 trends emerged across urban types: negative trends at a residential-industrial site, positive trends at a site surrounded by rapid suburban growth, and relatively constant CO2 over time at multiple sites in the established, residential, and commercial urban core. Analysis of population within the atmospheric footprints of the different sites reveals approximately equal increases in population influencing the observed CO2, implying a nonlinear relationship with CO2 emissions: Population growth in rural areas that experienced suburban development was associated with increasing emissions while population growth in the developed urban core was associated with stable emissions. Four state-of-the-art global-scale emission inventories also have a nonlinear relationship with population density across the city; however, in contrast to our observations, they all have nearly constant emissions over time. Our results indicate that decadal scale changes in urban CO2 emissions are detectable through monitoring networks and constitute a valuable approach to evaluate emission inventories and studies of urban carbon cycles.

A predictive spatial model for roasted coffee using oxygen isotopes of α-cellulose.

RATIONALE: Fraudulent region-of-origin labeling is a concern for high-value, globally traded commodities such as coffee. The oxygen isotope ratio of cellulose is a useful geographic tracer, as it integrates climate and source water signals. A predictive spatial model ("isoscape") of the δ18 O values of coffee bean cellulose is generated to evaluate coffee region-of-origin claims. METHODS: The oxygen isotope ratio of α-cellulose extracted from roasted coffee beans was measured via high-temperature conversion elemental analyzer/isotope ratio mass spectrometry (TC-EA/IRMS) and used to calculate the δ18 O value of coffee bean water. The 18 O enrichment of coffee bean water relative to the δ18 O value of local precipitation was modeled as a function of local temperature and humidity. This function was incorporated into a mechanistic model of cellulose δ18 O values to predict the δ18 O values of coffee bean cellulose across coffee-producing regions globally. RESULTS: The δ18 O values of analyzed coffee bean cellulose ranged from approximately +22‰ to +42‰ (V-SMOW). As expected, coffees grown in the same region tended to have similar isotope ratios, and the δ18 O value of coffee bean cellulose was generally higher than the δ18 O value of modeled stem cellulose for the region. Modeled δ18 O values of coffee cellulose were within ±2.3‰ of the measured δ18 O value of coffee cellulose. CONCLUSIONS: The oxygen isotope ratio of coffee bean cellulose is a useful indicator of region-of-origin and varies predictably in response to climatic factors and precipitation isotope ratios. The isoscape of coffee bean cellulose δ18 O values from this study provides a quantitative tool that can be applied to region-of-origin verification of roasted coffee at the point-of-sale.

Recent increases in drought frequency cause observed multi-year drought legacies in the tree rings of semi-arid forests.

Recent analyses on the length of drought recovery in forests have shown multi-year legacies, particularly in semi-arid, coniferous ecosystems. Such legacies are usually attributed to ecophysiological memory, although drought frequency itself, and its effect on overlapping recovery times, could also contribute. Here, we describe a multi-decadal study of drought legacies using tree-ring carbon-isotope ratios (δ13C) and ring-width index (RWI) in Pinus ponderosa at 13 montane sites traversing a winter-summer precipitation gradient in the Southwestern U.S. Sites and trees were selected to avoid collection biases that exist in archived tree-ring databanks. The spatial hydroclimate gradient and winter-summer seasonal patterns were well predicted by seasonal and inter-annual correlations between δ13C and atmospheric vapor pressure deficit (VPD). Using VPD, we found that the probability of extreme drought has increased up to 70% in this region during the past two decades. When the recent increase in drought frequency was not considered, multi-year legacies in both δ13C and RWI were observed at most sites. When the increase in drought frequency was detrended from tree-ring chronologies, some sites exhibited short legacies (1-2 years) in both δ13C and RWI, and there was a sight trend for longer legacies in RWI. However, when considered broadly across the region and multiple decades, no significant legacies were observed, which contrasts with past studies. Our results reveal that a contribution to observed multi-year legacies is related to shifts in the climate system itself, an exogenous factor, that must be considered along with physiological memory.

Climate and lawn management interact to control C4 plant distribution in residential lawns across seven U.S. cities.

In natural grasslands, C4 plant dominance increases with growing season temperatures and reflects distinct differences in plant growth rates and water use efficiencies of C3 vs. C4 photosynthetic pathways. However, in lawns, management decisions influence interactions between planted turfgrass and weed species, leading to some uncertainty about the degree of human vs. climatic controls on lawn species distributions. We measured herbaceous plant carbon isotope ratios (δ13 C, index of C3 /C4 relative abundance) and C4 cover in residential lawns across seven U.S. cities to determine how climate, lawn plant management, or interactions between climate and plant management influenced C4 lawn cover. We also calculated theoretical C4 carbon gain predicted by a plant physiological model as an index of expected C4 cover due to growing season climatic conditions in each city. Contrary to theoretical predictions, plant δ13 C and C4 cover in urban lawns were more strongly related to mean annual temperature than to growing season temperature. Wintertime temperatures influenced the distribution of C4 lawn turf plants, contrary to natural ecosystems where growing season temperatures primarily drive C4 distributions. C4 cover in lawns was greatest in the three warmest cities, due to an interaction between climate and homeowner plant management (e.g., planting C4 turf species) in these cities. The proportion of C4 lawn species was similar to the proportion of C4 species in the regional grass flora. However, the majority of C4 species were nonnative turf grasses, and not of regional origin. While temperature was a strong control on lawn species composition across the United States, cities differed as to whether these patterns were driven by cultivated lawn grasses vs. weedy species. In some cities, biotic interactions with weedy plants appeared to dominate, while in other cities, C4 plants were predominantly imported and cultivated. Elevated CO2 and temperature in cities can influence C3 /C4 competitive outcomes; however, this study provides evidence that climate and plant management dynamics influence biogeography and ecology of C3 /C4 plants in lawns. Their differing water and nutrient use efficiency may have substantial impacts on carbon, water, energy, and nutrient budgets across cities.

Strontium isotope ratios of human hair from the United States: Patterns and aberrations.

RATIONALE: Strontium isotope ratios (87 Sr/86 Sr) of hair may be a valuable tool to estimate human provenance. However, the systematics and mechanisms controlling spatial variation in 87 Sr/86 Sr of modern human hair remain unclear. Here, we measure 87 Sr/86 Sr of hair specimens from across the USA to assess the presence of geospatial relationships. METHODS: Ninety-eight human hair specimens were collected from salon/barbershop floors in 48 municipalities throughout the conterminous USA. [Sr] and 87 Sr/86 Sr ratios were measured from hair using quadrupole and multi-collector inductively coupled plasma mass spectrometers, respectively. The [Sr] and 87 Sr/86 Sr ratios of hair were compared with the measured [Sr] and 87 Sr/86 Sr ratios of tap waters from the collection locations. In addition, the 87 Sr/86 Sr ratio of hair was compared with the modeled ratios of bedrock and surface waters. RESULTS: Hair color was independent of the 87 Sr/86 Sr ratio, but related to [Sr]. The 87 Sr/86 Sr ratios of hair and leachate were not statistically different and were positively correlated; however, in several hair-leachate pairs, the ratios were conspicuously different. The 87 Sr/86 Sr ratios of both hair and leachate were linearly correlated with tap water. The 87 Sr/86 Sr ratio of hair was also significantly correlated with the modeled ratio of bedrock and surface waters, although the 87 Sr/86 Sr ratio of hair was most strongly correlated with the measured ratio of tap water. CONCLUSIONS: The 87 Sr/86 Sr ratio of hair is related to the ratio of tap water, which varied geographically. The ratio of hair provided geographic information about an individual's recent residence. Differences in the 87 Sr/86 Sr ratios of hair and hair leachate may be concomitant with travel and could potentially be used as a screening tool to identify recent movements.

Disentangling seasonal and interannual legacies from inferred patterns of forest water and carbon cycling using tree-ring stable isotopes.

Tree-ring carbon and oxygen isotope ratios have been used to understand past dynamics in forest carbon and water cycling. Recently, this has been possible for different parts of single growing seasons by isolating anatomical sections within individual annual rings. Uncertainties in this approach are associated with correlated climate legacies that can occur at a higher frequency, such as across successive seasons, or a lower frequency, such as across years. The objective of this study was to gain insight into how legacies affect cross-correlation in the δ13 C and δ18 O isotope ratios in the earlywood (EW) and latewood (LW) fractions of Pinus ponderosa trees at thirteen sites across a latitudinal gradient influenced by the North American Monsoon (NAM) climate system. We observed that δ13 C from EW and LW has significant positive cross-correlations at most sites, whereas EW and LW δ18 O values were cross-correlated at about half the sites. Using combined statistical and mechanistic models, we show that cross-correlations in both δ13 C and δ18 O can be largely explained by a low-frequency (multiple-year) mode that may be associated with long-term climate change. We isolated, and statistically removed, the low-frequency correlation, which resulted in greater geographical differentiation of the EW and LW isotope signals. The remaining higher-frequency (seasonal) cross-correlations between EW and LW isotope ratios were explored using a mechanistic isotope fractionation-climate model. This showed that lower atmospheric vapor pressure deficits associated with monsoon rain increase the EW-LW differentiation for both δ13 C and δ18 O at southern sites, compared to northern sites. Our results support the hypothesis that dominantly unimodal precipitation regimes, such as near the northern boundary of the NAM, are more likely to foster cross-correlations in the isotope signals of EW and LW, potentially due to greater sharing of common carbohydrate and soil water resource pools, compared to southerly sites with bimodal precipitation regimes.

Strontium isotope ratios (87 Sr/86 Sr) of human fingernail clippings reveal multiple location signals.

RATIONALE: Strontium isotope ratios (87 Sr/86 Sr) in human fingernail keratin tissues have been underexplored for region of origin and travel history reconstruction studies. Here we investigated 87 Sr/86 Sr ratios in fingernail keratin to establish baseline measurements in a resident group and to examine how 87 Sr/86 Sr ratios changed with relocation. METHODS: Fingernail clippings were collected from resident (n = 10) and non-resident/traveler groups (n = 4 and n = 4) that were part of a larger study in Salt Lake City (UT, USA) from 2015 to 2016. Strontium abundance and 87 Sr/86 Sr ratios were determined via multicollector inductively coupled plasma mass spectrometry (MC-ICP-MS). 87 Sr/86 Sr and oxygen (δ18 O) isotope ratios from the traveler participants were compared to examine temporal patterns. RESULTS: Strontium abundance and 87 Sr/86 Sr ratios in fingernails from the resident group established a baseline against which we could evaluate potential differences in non-resident/traveler groups. Resident 87 Sr/86 Sr ratios remained constant over the study period and were consistent with previously measured tap waters for the area. 87 Sr/86 Sr ratio changes in non-resident/traveler groups were rapid and reflected the current location of the individual within 4-5 weeks of arrival. Lastly, δ18 O and 87 Sr/86 Sr ratios of the same fingernail clippings did not exhibit similar temporal patterns, since fingernail δ18 O ratios required more time to attain values characteristic of the new environment. CONCLUSIONS: Our findings suggest that strontium is incorporated into fingernail tissues differently from hair and this could be advantageous to forensic investigations. We found that 87 Sr/86 Sr and δ18 O ratios of the same fingernail clippings revealed two different time points reflecting an individual's residence over short- (4-5 weeks, 87 Sr/86 Sr ratios) and long-term (3-5 months, δ18 O values) time scales. It is likely that the 87 Sr/86 Sr ratios of fingernail clippings reflect exogenous signals that are incorporated through bathing waters and that these signals change rapidly with movement to a new location. Our results may aid future forensic studies in the determination of region of origin in unidentified remains.

Distinctions in heterotrophic and autotrophic-based metabolism as recorded in the hydrogen and carbon isotope ratios of normal alkanes.

The hydrogen isotope values of n-alkanes (δ2Hn-alkane) reflect a plant's water source and water relations, while the carbon isotope values (δ13Cn-alkane) relate to a plant's carbon metabolism and response to environmental conditions. However, the isotopic dynamics of the transition from heterotrophic to autotrophic metabolism during foliar development on δ2Hn-alkane and δ13Cn-alkane remain unclear. Here, we monitored δ2Hn-alkane and δ13Cn-alkane across a growing season from Betula occidentalis, Populus angustifolia, and Acer negundo. In addition, we compiled δ2H values of atmospheric vapor, leaf water, xylem water, and stream water as well as δ13C values of bulk leaf tissue (δ13Cbulk). We found δ2Hn-alkane and δ13Cn-alkane varied with leaf development and indicated that the majority of wax development occurred during the initial growing season. The patterns in δ2Hn-alkane were broadly consistent between species and with previous studies; however, each species had a unique final δ2Hn-alkane value. The δ13Cbulk for all species demonstrated a characteristic 13C-enrichment during the initial growing season, followed by 13C-depletion, while δ13Cn-alkane did not exhibit a consistent trend between the species. These δ13C data suggested a decoupling of the isotopic inputs between n-alkanes and photosynthetic leaf tissue. When coupled with δ2Hn-alkane, these data suggested that the precursor compounds utilized in initial production of n-alkanes might be variable and possibly indicated that the stored precursors used for initial leaf tissue and wax production originated from different sources. Nonetheless, these data indicated that the isotopic signatures of n-alkanes relate to a mixture of precursors, but only during a distinct period of leaf ontogeny.

A tale of ENSO, PDO, and increasing aridity impacts on drought-deciduous shrubs in the Death Valley region.

Germination, establishment, phenology, and death among three drought-deciduous shrubs were influenced by ENSO/PDO and precipitation, based on 37 years of annual surveys. Encelia farinosa forms near monospecific stands on slopes, whereas E. frutescens and Ambrosia salsola dominate wash habitats. All shrubs exhibited phenological coherence. While germination, establishment, and mortality patterns were similar among wash species, these dynamics contrasted with E. farinosa on slopes. Germination was associated with El Niño years. Slope plant establishment was dependent on precipitation in the subsequent year, but not evidently so in wash species. Major mortality events were episodic, with Encelia mortality just as likely to occur in years with below or above average precipitation. In both Encelia species, mortality was associated with transitions to a cold PDO phase. In E. frutescens this response was more rapid, whereas in E. farinosa mortality lagged 1 year, resulting in contrasting slope-wash mortality patterns. 50% of newly established shrubs died within 5, 5, and 18 years for E. frutescens, E. farinosa, and A. salsola, respectively. The 90% mortality ages were 26 years for E. frutescens, 24 years for E. farinosa, and 51 years for A. salsola. While maximum life expectancies are unknown, estimated maximum life expectancies were 56, 66, and 86 years for E. frutescens, E. farinosa, and A. salsola, respectively. Overall, as the climate has become more arid over the past four decades, the populations in both slope and wash habitats have exhibited similar responses: reduced shrub abundances and reduced total supportable leaf areas.

Some like it hot: the physiological ecology of C4 plant evolution.

The evolution of C4 photosynthesis requires an intermediate phase where photorespiratory glycine produced in the mesophyll cells must flow to the vascular sheath cells for metabolism by glycine decarboxylase. This glycine flux concentrates photorespired CO2 within the sheath cells, allowing it to be efficiently refixed by sheath Rubisco. A modest C4 biochemical cycle is then upregulated, possibly to support the refixation of photorespired ammonia in sheath cells, with subsequent increases in C4 metabolism providing incremental benefits until an optimized C4 pathway is established. 'Why' C4 photosynthesis evolved is largely explained by ancestral C3 species exploiting photorespiratory CO2 to improve carbon gain and thus enhance fitness. While photorespiration depresses C3 performance, it produces a resource (photorespired CO2) that can be exploited to build an evolutionary bridge to C4 photosynthesis. 'Where' C4 evolved is indicated by the habitat of species branching near C3-to-C4 transitions on phylogenetic trees. Consistent with the photorespiratory bridge hypothesis, transitional species show that the large majority of > 60 C4 lineages arose in hot, dry, and/or saline regions where photorespiratory potential is high. 'When' C4 evolved has been clarified by molecular clock analyses using phylogenetic data, coupled with isotopic signatures from fossils. Nearly all C4 lineages arose after 25 Ma when atmospheric CO2 levels had fallen to near current values. This reduction in CO2, coupled with persistent high temperature at low-to-mid-latitudes, met a precondition where photorespiration was elevated, thus facilitating the evolutionary selection pressure that led to C4 photosynthesis.