Tree rings are considered the gold standard for observing variation in past atmospheric radiocarbon (14C), yet little attention has been paid to whether different trees record tropospheric 14C evenly. The discovery of Miyake events, rapid increases in 14C production occurring in past millennia, has led to repeated measurements of 14C in tree rings across species and locations. These records demonstrate remarkable synchrony, yet significant variability between samples remains, limiting reliable use of tree rings as a precise indicator of the timing and scale of past 14C production. Understanding how trees and species record tropospheric 14C would improve reconstructions of past 14C production events, refine geochronological control, and because of the pulse-like nature of past 14C production events, shed light on a fundamental question in tree physiology and dendrochronology: what is the lag time between photosynthesis, storage, and allocation to wood and how might this lag blur isotopic signals in tree rings? Here, we evaluate the implicit assumption that all trees record tropospheric 14C evenly by focusing on the path carbon takes within trees including: phenology of carbon uptake and wood formation, storage and use of nonstructural carbohydrates, and how these factors interact to affect the tropospheric 14C signal in wood.

In forests of the southwestern USA, the seasonality-or phenology-of tree growth is affected by a combination of limiting temperatures and water availability. However, in this topographically diverse area, temperature and precipitation vary by elevation and therefore may have differing effects on tree phenology across the landscape. Our over-arching research question was: how does variation in temperature and water availability drive differences in the timing of tree growth and water use across species and sites in northern Arizona? We analyze 3 years of high-frequency measurements of stem growth and sap flow velocity collected across five sites from 1400 to 2600 meters elevation. We supplemented these data with lower-frequency measurements of xylogenesis, budburst, and potential photosynthetic quantum efficiency. Our study species-Pinus ponderosa, Pinus edulis, Juniperus osteosperma, Juniperus scopulorum and Quercus gambelii-represent the dominant tree species across northern Arizona. We found that all metrics of tree phenology tended to be limited by water availability at lower elevations. Lower elevations had bimodal patterns of growth and sap flow velocity which reflected precipitation inputs. Higher elevations had more consistent unimodal patterns which aligned with daylength, and growth was rarely limited by water availability. Microcore data supported growth rates from dendrometers, but microcores were able to capture xylogenesis when dendrometers were unable to-surprisingly, even when stem diameter was shrinking due to water limitation. A machine learning model showed soil volumetric water content was the best predictor of radial growth and sap flow velocity at lower elevations but was only marginally better than daylength and temperature at higher elevations. Thus, tree activity in trailing-edge, low elevation forests was more impacted by moisture, and less affected by temperature, compared with forests at higher elevations. Our results, which show that arid woodlands and forests in the North American Southwest are adapted to grow opportunistically when neither water availability nor air temperature is limiting, are synthesized in a new conceptual model.

Constrained carbon allocation toward secondary metabolites involved in chemical defense is a common explanation for widespread drought-related beetle-kill in conifers-we challenge the generality of this explanation. While monitoring drought stress (ψpd), we tracked both carbon reserves (non-structural carbohydrates) and chemical defenses (terpenes, phenolics, resin flow) in mature Pinus edulis Englem. trees experiencing either short-term (3-year) or a 'legacy' long-term (13-year) throughfall exclusion treatments, plus a control. We also quantified the Δ14C-age of resin to measure past allocation to current defense. While 72% of trees in short-term throughfall exclusion plots died (attacked by bark beetles, Ips confusus LeConte), mortality patterns were unrelated to throughfall exclusion intensity and all 'legacy' trees survived. We thus assessed trees in four survivorship categories: control, 'legacy', surviving, and dying trees. We found concentrations of certain defense compounds (leaf phenolics, twig monoterpenes) increased with drought stress, particularly in dying trees. In the main stem, dying trees exhibited similar terpene concentrations (94%) and phenolic concentrations (139%) relative to control trees. Compared with control trees, only 'legacy' trees had reduced stem terpenes (-49%, P < 0.05) after a decade of drought. Δ14C-age of resin could be up to 10.2 ± 0.5 years old, where the oldest resin was exuded from trees with low sugar concentrations and more negative Ψpd. Our results suggest that drought imposes a weak constraint on carbon allocation to resin-based defense. Instead, we primarily found evidence of increased concentrations of terpene and phenolic compounds under drought, even in dying trees, and only observed reductions in resin-based defenses after 10+ years of drought. Δ14C-ages demonstrate limited resin turnover and/or synthesis of resin from old reserves, suggesting that long-term drought is required to reduce resin-based defenses. Persistent allocation coupled with past investments appears to preserve or enhance concentrations of resin-based defenses even under lethal drought stress in P. edulis.

A declining spring snowpack is expected to have widespread effects on montane and subalpine forests in western North America and across the globe. The way that tree water demands respond to this change will have important impacts on forest health and downstream water subsidies. Here, we present data from a network of sap velocity sensors and xylem water isotope measurements from three common tree species (Picea engelmannii, Abies lasiocarpa and Populus tremuloides) across a hillslope transect in a subalpine watershed in the Upper Colorado River basin. We use these data to compare tree- and stand-level responses to the historically high spring snowpack but low summer rainfall of 2019 against the low spring snowpack but high summer rainfall amounts of 2021 and 2022. From the sap velocity data, we found that only 40 % of the trees showed an increase in cumulative transpiration in response to the large snowpack year (2019), illustrating the absence of a common response to interannual spring snowpack variability. The trees that increased water use during the year with the large spring snowpack were all found in dense canopy stands – irrespective of species – while trees in open-canopy stands were more reliant on summer rains and, thus, more active during the years with modest snow and higher summer rain amounts. Using the sap velocity data along with supporting measurements of soil moisture and snow depth, we propose three mechanisms that lead to stand density modulating the tree-level response to changing seasonality of precipitation: Topographically mediated convergence zones have consistent access to recharge from snowmelt which supports denser stands with high water demands that are more reliant and sensitive to changing snow. Interception of summer rain in dense stands reduces the throughfall of summer rain to surface soils, limiting the sensitivity of the dense stands to changes in summer rain. Shading in dense stands allows the snowpack to persist deeper into the growing season, providing high local reliance on snow during the fore-summer (early-summer) drought period. Combining data generated from natural gradients in stand density, like this experiment, with results from controlled forest-thinning experiments can be used to develop a better understanding of the responses of forested ecosystems to futures with reduced spring snowpack.

Soil respiration (SR) is a key component of terrestrial carbon-climate feedbacks, yet its seasonal dynamics in drylands remain poorly understood. In mesic ecosystems, SR is primarily temperature driven, whereas in dry-lands it shifts seasonally from temperature to moisture control as autotrophic and heterotrophic respiration become water limited during dry periods. Identifying the soil temperature at which SR transitions from temperature to moisture limitation is therefore essential for predicting SR under climate change. We examined temperature and moisture response functions of SR across forests, shrublands, and grasslands in arid and semiarid regions to determine the soil temperature threshold of SR (STTSR) and its drivers. Across sites, SR was positively correlated with mean annual precipitation, soil moisture, and soil organic carbon, while negatively correlated with soil temperature. The significant variability in the temperature thresholds of SR (STTSR) that was observed between sites (17.9 degrees C f 5.3 degrees C; mean f SD) was best explained by the mean annual temperature (MAT) at the site. Sites with higher air temperatures exhibited higher STTSR, suggesting that the compartments and metabolic processes involved in SR are adapted to local temperatures. This observed SR adaptation occurred at two different scales. Besides STTSR were positively correlated with MAT within each vegetation type, STTSR were systematically higher under short-stature vegetation types (grasslands and shrublands) compared to high-stature vegetation types (forests), suggesting that grasses and shrubs have developed the evolutionary capacity to push the STTSR to warmer temperatures and hence withstand better drought stress than trees. Our findings suggest that: (1) process-based models assuming simple linear or exponential SR-temperature relationships overestimate SR in water-limited ecosystems; and (2) projected warming and increasing water scarcity, together with shifts in vegetation dominance, may strongly modify the temperature sensitivity of SR.

Carbon reserves are distributed throughout plant cells allowing past photosynthesis to fuel current metabolism. In trees, comparing the radiocarbon (Δ14C) of reserves to the atmospheric bomb spike can trace reserve ages. We synthesized Δ14C observations of stem reserves in nine tree species, fitting a new process model of reserve building. We asked how the distribution, mixing, and turnover of reserves vary across trees and species. We also explored how stress (drought and aridity) and disturbance (fire and bark beetles) perturb reserves. Given sufficient sapwood, young (< 1 yr) and old (20-60+ yr) reserves were simultaneously present in single trees, including 'prebomb' reserves in two conifers. The process model suggested that most reserves are deeply mixed (30.2 ± 21.7 rings) and then respired (2.7 ± 3.5-yr turnover time). Disturbance strongly increased Δ14C mean ages of reserves (+15-35 yr), while drought and aridity effects on mixing and turnover were species-dependent. Fire recovery in Sequoia sempervirens also appears to involve previously unobserved outward mixing of old reserves. Deep mixing and rapid turnover indicate most photosynthate is rapidly metabolized. Yet ecological variation in reserve ages is enormous, perhaps driven by stress and disturbance. Across species, maximum reserve ages appear primarily constrained by sapwood longevity, and thus old reserves are probably widespread.

Long-term soil CO _2 emission measurements are necessary for detecting trends and interannual variability in the terrestrial carbon cycle. Such records are becoming increasingly valuable as ecosystems experience altered environmental conditions associated with climate change. From 2013 to 2021, we continuously measured soil CO _2 concentrations in the two dominant high elevation forest types, mixed conifer and aspen, in the upper Colorado River basin. We quantified the soil CO _2 flux during the summer months, and found that the mean and total CO _2 flux in both forests was related to the prior winter’s snowfall and current summer’s rainfall, with greater sensitivity to rainfall. We observed a decline in surface soil CO _2 production, which we attributed to warming and a decrease in amount and frequency of summer rains. Our results demonstrate strong precipitation control on the soil CO _2 flux in mountainous regions, a finding which has important implications for carbon cycling under future environmental change.

Radiocarbon (∆14C) measurements of nonstructural carbon enable inference on the age and turnover time of stored photosynthate (e.g., sugars, starch), of which the largest pool in trees resides in the main bole. Because of potential issues with extraction-based methods, we introduce an incubation method to capture the ∆14C of nonstructural carbon via respired CO2. In this study, we compared the ∆14C obtained from these incubations with ∆14C from a well-established extraction method, using increment cores from a mature trembling aspen (Populus tremuloides). To understand any potential ∆14C disagreement, the yields from both methods were also benchmarked against the phenol-sulfuric acid concentration assay. We found incubations captured less than 100% of measured sugar and starch carbon, with recovery ranging from ~ 3% in heartwood to 85% in shallow sapwood. However, extractions universally over-yielded (mean 273 ± 101% expected sugar carbon; as high as 480%), where sugars represented less than half of extracted soluble carbon, indicating very poor specificity. While separation of soluble and insoluble nonstructural carbon is ostensibly a strength of extraction based methods, there was also evidence of poor separation of these two fractions in extractions. The ∆14C of respired CO2 and ∆14C from extractions were similar in the sapwood, while extractions resulted in comparatively higher ∆14C (older carbon) in heartwood and bark. Because yield and ∆14C discrepancies were largest in old tissues, incubations may better capture the ∆14C of nonstructural carbon that is actually metabolically available. That is, we suggest extractions include metabolically irrelevant carbon from dead tissues or cells, as well as carbon that is neither sugar nor starch. In contrast, nonstructural carbon captured by extractions must be respired to be measured. We thus suggest incubations of live tissues are a potentially viable, inexpensive, and versatile method to study the ∆14C of metabolically relevant (available) nonstructural carbon.

ABSTRACTWe present a timeseries of 14CO2 for the period 1910–2021 recorded by annual plants collected in the southwestern United States, centered near Flagstaff, Arizona. This timeseries is dominated by five commonly occurring annual plant species in the region, which is considered broadly representative of the southern Colorado Plateau. Most samples (1910–2015) were previously archived herbarium specimens, with additional samples harvested from field experiments in 2015–2021. We used this novel timeseries to develop a smoothed local record with uncertainties for “bomb spike” 14C dating of recent terrestrial organic matter. Our results highlight the potential importance of local records, as we document a delayed arrival of the 1963–1964 bomb spike peak, lower values in the 1980s, and elevated values in the last decade in comparison to the most current Northern Hemisphere Zone 2 record. It is impossible to retroactively collect atmospheric samples, but archived annual plants serve as faithful scribes: samples from herbaria around the Earth may be an under-utilized resource to improve understanding of the modern carbon cycle.

The Greenness_Snodgrass data set is downloaded and reorganized data from the PhenoCam website. All Phenocam metric column headers are unchanged from the imported PhenoCam server. The data was downloaded using Rstudio. And package phenocamr. This raw downloaded data is described in detail in Richardson, A. D. et al. Tracking vegetation phenology across diverse North American biomes using PhenoCam imagery. Sci. Data 5:180028 doi: 10.1038/sdata.2018.28 (2018). This data set has taken the above-described data and merged data record 4 and data record 5 at each site on the elevation transect. Added columns that are not described in Richardson et al 2019 include species (colloquial plant type), site(abbreviated Snodgrass site name ), and transition_type (classifies the transition date as falling or rising).

Shifts in the age or turnover time of non-structural carbohydrates (NSC) may underlie changes in tree growth under long-term increases in drought stress associated with climate change. But NSC responses to drought are challenging to quantify, due in part to large NSC stores in trees and subsequently long response times of NSC to climate variation. We measured NSC age (Δ14 C) along with a suite of ecophysiological metrics in Pinus edulis trees experiencing either extreme short-term drought (-90% ambient precipitation plot, 2020-2021) or a decade of severe drought (-45% plot, 2010-2021). We tested the hypothesis that carbon starvation - consumption exceeding synthesis and storage - increases the age of sapwood NSC. One year of extreme drought had no impact on NSC pool size or age, despite significant reductions in predawn water potential, photosynthetic rates/capacity, and twig and needle growth. By contrast, long-term drought halved the age of the sapwood NSC pool, coupled with reductions in sapwood starch concentrations (-75%), basal area increment (-39%), and bole respiration rates (-28%). Our results suggest carbon starvation takes time, as tree carbon reserves appear resilient to extreme disturbance in the short term. However, after a decade of drought, trees apparently consumed old stored NSC to support metabolism.

The data set “Snowpack_Snodgrass_AS.csv” consists of the snowpack data measured by the PhenoCams at each site up Snodgrass Mtn, Crested Butte, Co. To calculate snow depth each fall, we placed a 3-meter PVC pipe in the ground in the field of view of the PhenoCam. We spray painted 25 cm alternating increments of red and white, with each meter labeled with a black stripe. Using the downloaded PhenoCam images, a calibration curve of pole height to pixels was calculated. The amount of non-snow-covered pole pixels for each image was quantified, and the snow depth was calculated using the calibration curve. Sites are abbreviated versions of site names and phenocam numbers, i.e., hc7= high_conifer_Snodgrass_7.

For long-lived organisms, investment in insurance strategies such as reserve energy storage can enable resilience to resource deficits, stress or catastrophic disturbance. Recent fire in California damaged coast redwood (Sequoia sempervirens) groves, consuming all foliage on some of the tallest and oldest trees on Earth. Burned trees recovered through resprouting from roots, trunk and branches, necessarily supported by nonstructural carbon reserves. Nonstructural carbon reserves can be many years old, but direct use of old carbon has rarely been documented and never in such large, old trees. We found some sprouts contained the oldest carbon ever observed to be remobilized for growth. For certain trees, simulations estimate up to half of sprout carbon was acquired in photosynthesis more than 57 years prior, and direct observations in sapwood show trees can access reserves at least as old. Sprouts also emerged from ancient buds-dormant under bark for centuries. For organisms with millennial lifespans, traits enabling survival of infrequent but catastrophic events may represent an important energy sink. Remobilization of decades-old photosynthate after disturbance demonstrates substantial amounts of nonstructural carbon within ancient trees cycles on slow, multidecadal timescales. 14C measurements show coast redwoods (Sequoia sempervirens) remobilize decades- to century-old carbon reserves to regrow leaves after a rare fire. To do so, trees resprout from dormant bud tissue maintained for centuries or more.

Linking biometric measurements of stand‐level biomass growth to tower‐based measurements of carbon uptake—gross primary productivity and net ecosystem productivity—has been the focus of numerous ecosystem‐level studies aimed to better understand the factors regulating carbon allocation to slow‐turnover wood biomass pools. However, few of these studies have investigated the importance of previous year uptake to growth. We tested the relationship between wood biomass increment (WBI) and different temporal periods of carbon uptake from the current and previous years to investigate the potential lagged allocation of fixed carbon to growth among six mature, temperate forests. We found WBI was strongly correlated to carbon uptake across space (i.e., long‐term averages at the different sites) but on annual timescales, WBI was much less related to carbon uptake, suggesting a temporal mismatch between C fixation and allocation to biomass. We detected lags in allocation of the previous year's carbon uptake to WBI at three of the six sites. Sites with higher annual WBI had overall stronger correlations to carbon uptake, with the strongest correlations to carbon uptake from the previous year. Only one site had WBI with strong positive relationships to current year uptake and not the previous year. Forests with low rates of WBI demonstrated weak correlations to carbon uptake from the previous year and stronger relationships to current year climate conditions. Our work shows an important, but not universal, role of lagged allocation of the previous year's carbon uptake to growth in temperate forests.

Biological soil crusts (biocrusts) are critical components of dryland and other ecosystems worldwide, and are increasingly recognized as novel model ecosystems from which more general principles of ecology can be elucidated. Biocrusts are often diverse communities, comprised of both eukaryotic and prokaryotic organisms with a range of metabolic lifestyles that enable the fixation of atmospheric carbon and nitrogen. However, how the function of these biocrust communities varies with succession is incompletely characterized, especially in comparison to more familiar terrestrial ecosystem types such as forests. We conducted a greenhouse experiment to investigate how community composition and soil-atmosphere trace gas fluxes of CO2, CH4, and N2O varied from early-successional light cyanobacterial biocrusts to mid-successional dark cyanobacteria biocrusts and late-successional moss-lichen biocrusts and as biocrusts of each successional stage matured. Cover type richness increased as biocrusts developed, and richness was generally highest in the late-successional moss-lichen biocrusts. Microbial community composition varied in relation to successional stage, but microbial diversity did not differ significantly among stages. Net photosynthetic uptake of CO2 by each biocrust type also increased as biocrusts developed but tended to be moderately greater (by up to ≈25%) for the mid-successional dark cyanobacteria biocrusts than the light cyanobacterial biocrusts or the moss-lichen biocrusts. Rates of soil C accumulation were highest for the dark cyanobacteria biocrusts and light cyanobacteria biocrusts, and lowest for the moss-lichen biocrusts and bare soil controls. Biocrust CH4 and N2O fluxes were not consistently distinguishable from the same fluxes measured from bare soil controls; the measured rates were also substantially lower than have been reported in previous biocrust studies. Our experiment, which uniquely used greenhouse-grown biocrusts to manipulate community composition and accelerate biocrust development, shows how biocrust function varies along a dynamic gradient of biocrust successional stages.

1.ABSTRACTBackgroundDetermining the life-history traits of extinct species is often difficult from skeletal remains alone, limiting the accuracy of studies modeling past ecosystems. However, the analysis of the degraded endogenous bacterial DNA present in paleontological fecal matter (coprolites) may enable the characterization of specific traits such as the host’s digestive physiology and diet. An issue when evaluating the microbial composition of coprolites is the degree to which the microbiome is representative of the host’s original gut community versus the changes that occur in the weeks following deposition due to desiccation. Analyses of paleontological microorganisms are also relevant in the light of recent studies linking the Late Pleistocene and Early Holocene extinctions with modern-day zoonotic pathogen outbreaks.MethodsShotgun sequencing was performed on ancient DNA (aDNA) extracted from coprolites of the Columbian mammoth (Mammuthus Columbi), Shasta ground sloth (Nothrotheriops shastensis) and paleontological bison (Bison sp.) collected from caves on the Colorado Plateau, Southwestern USA. The novel metagenomic classifier MTSv, parameterized for studies of aDNA, was used to assign bacterial taxa to sequencing reads. The resulting bacterial community of coprolites was then compared to those from modern fecal specimens of the African savannah elephant (Loxodonta africana), the brown-throated sloth (Bradypus variegatus) and the modern bison (Bison bison). Both paleontological and modern bison fecal bacterial communities were also compared to those of progressively dried cattle feces to determine whether endogenous DNA from coprolites had a microbiome signal skewed towards aerobic microorganisms typical of desiccated fecal matter.ResultsThe diversity of phyla identified from coprolites was lower than modern specimens. The relative abundance of Actinobacteria was increased in coprolites compared to modern specimens, with fewer Bacteroidetes and Euryarchaeota. Firmicutes had a reduced relative abundance in the mammoth and bison coprolites, compared to the African savanna elephants and modern bison. There was a significant separation of samples in NMDS plots based on their classification as either paleontological or modern, and to a lesser extent, based on the host species. Increasingly dried cattle feces formed a continuum between the modern and paleontological bison samples.ConclusionOur results reveal that any coprolite metagenomes should always be compared to desiccated modern fecal samples from closely related hosts fed a comparable diet to determine the degree to which the coprolite metagenome is a result of desiccation versus true dissimilarities between the modern and paleontological hosts. Also, a large-scale desiccation study including a variety of modern species may shed light on life-history traits of extinct species without close extant relatives, by establishing the proximity of coprolite metagenomes with those from dried modern samples.

Under increasingly frequent, persistent, and severe drought events, predicting future forest carbon dynamics necessitates quantitative understanding of the physiological processes leading to tree mortality and physiological impairment. The responses of non-structural carbon (NSC; primarily sugars and starch) pools in mature trees is particularly important, as dynamics in NSC interact with hydraulic damage to perturb future tree growth. However, NSC concentration measurements alone are not suUcient to understand the stress responses of tree NSC pools formed over years to decades. Thus, we are using radiocarbon (14C) to quantify the age of NSC stored within, and used by, piñon pine trees exposed to either severe or long-term drought stress at the Sevilleta LTER, in New Mexico, USA. Measuring the age of NSC allows inference on the storage history of a tree, and how different NSC pools may be altered by drought. Experimental plots are subjected to either 0% (control) or 90% reduction in precipitation. A 45% precipitation reduction plot has also been in place since 2009, offering a chance to study the impacts of a decade of drought. We are measuring Δ14C of NSC in twigs, bole sapwood, and coarse roots, as well as in CO2 respired from the bole and branches. Our goal is to quantify the role of different-aged NSC pools across tree organs in driving whole-tree physiological responses to drought. Preliminary results show that the long-term droughted trees store and respire on average younger NSC than control trees. Ongoing drought treatments and sampling will provide additional information on how NSC dynamics in these trees are influenced by drought.

The selective use of seasonal precipitation by vegetation is critical to understanding the residence time and flow path of water in watersheds, yet there are limited datasets to test how climate alters these dynamics. Here, we use measurements of the seasonal cycle of tree ring delta 18O for two widespread conifer species in the Rocky Mountains of North America to provide a multi-decadal depiction of the seasonal origins of forest water use. The results show that while the conifer tree stands had a dominant preference for use of snowmelt, there were multi-annual periods over the last four decades when use of summer precipitation was preferential. Utilization of summer rain emerged during years with increased snowfall and tree growth, suggesting that summer rain enhanced the transpiration stream only during the periods of highest water use. We hypothesize this could be explained through shallowing of the root profile during wetter periods and/or through the influence of changing water table depths on the residence time of summer precipitation in the root zone. We suggest the tree ring proxy approach used here could be applied in other watersheds to provide critical insight into the temporal dynamics of plant water use that could not be inferred from short measurement campaigns. These data on the seasonal origins of forest water are critical for understanding forest vulnerability to drought, the processes that affect precipitation pathways and residence time in watersheds and the interpretation of tree ring proxy data. Key Points Cellulose isotope ratios were used to reconstruct the reliance on snowmelt for two common conifer species in the southern Rocky Mountains The trees exhibited multi-year periods of preferential use of snowmelt and low tree growth following low snowpack years During high snowpack periods, the trees showed enhanced growth and utilized summer precipitation to support their increased water demands