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International audience; Background/Question/Methods Tropical forest responses to moisture remain poorly understood, in part because of the large diversity of plant hydraulic traits found therein. Changing moisture regimes, such as more frequent drought events, are expected to interact with these diverse hydraulic traits and other requirements of tropical trees in complex ways, making prediction of ecosystem-scale responses and community compositional trajectories difficult. A first step towards discerning such responses is in the analysis of how plant hydraulic and edaphic conditions control trajectories of individual trees’ water use over pre-drought, drought, and recovery periods. We took advantage of the 2015-2016 ENSO event, which induced drought over much of the tropics, to collect sap flow data on 47 canopy and numerous sub-canopy trees across nine sites in Latin America varying in annual precipitation from 1700 mm to > 3000 mm. This enabled us to determine a range of responses to changes in moisture, both atmospheric (VPD) and in soil. Where available, via measurements on conspecific individuals or species-mean values in trait databases, plant hydraulic traits were associated with individual sap flux trajectories, in addition to site-specific soil properties and climate. Results/Conclusions We found a large heterogeneity of sap flow responses during the ENSO within and among study regions. The diversity of strategies to deal with drought stress was partially explained by species functional traits, background climate and intensity of soil water depletion during the ENSO. Preliminary simulations of drought responses using the Community Land Model coupled to the hydraulically-enabled Functionally Assembled Terrestrial Ecosystem Simulator (CLM-FATES-Hydro) were used to demonstrate multiple mechanisms, both edaphic- and plant trait-related, responsible for the divergence in observed sap flow responses, as well as highlight critical field measurements needed to discern among these mechanisms.

© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust The fate of tropical forests under climate change is unclear as a result, in part, of the uncertainty in projected changes in precipitation and in the ability of vegetation models to capture the effects of drought-induced mortality on aboveground biomass (AGB). We evaluated the ability of a terrestrial biosphere model with demography and hydrodynamics (Ecosystem Demography, ED2-hydro) to simulate AGB and mortality of four tropical tree plant functional types (PFTs) that operate along light- and water-use axes. Model predictions were compared with observations of canopy trees at Barro Colorado Island (BCI), Panama. We then assessed the implications of eight hypothetical precipitation scenarios, including increased annual precipitation, reduced inter-annual variation, El Niño-related droughts and drier wet or dry seasons, on AGB and functional diversity of the model forest. When forced with observed meteorology, ED2-hydro predictions capture multiple BCI benchmarks. ED2-hydro predicts that AGB will be sustained under lower rainfall via shifts in the functional composition of the forest, except under the drier dry-season scenario. These results support the hypothesis that inter-annual variation in mean and seasonal precipitation promotes the coexistence of functionally diverse PFTs because of the relative differences in mortality rates. If the hydroclimate becomes chronically drier or wetter, functional evenness related to drought tolerance may decline.

© 2018, Springer International Publishing AG, part of Springer Nature. Background and aims: Forest tree microbiomes are important to forest dynamics, diversity, and ecosystem processes. Mature limber pines (Pinus flexilis) host a core microbiome of acetic acid bacteria in their foliage, but the bacterial endophyte community structure, variation, and assembly across tree ontogeny is unknown. The aims of this study were to test if the core microbiome observed in adult P. flexilis is established at the seedling stage, if seedlings host different endophyte communities in root and shoot tissues, and how environmental factors structure seedling endophyte communities. Methods: The 16S rRNA gene was sequenced to characterize the bacterial endophyte communities in roots and shoots of P. flexilis seedlings grown in plots at three elevations at Niwot Ridge, Colorado, subjected to experimental treatments (watering and heating). The data was compared to previously sequenced endophyte communities from adult tree foliage sampled in the same year and location. Results: Seedling shoots hosted a different core microbiome than adult tree foliage and were dominated by a few OTUs in the family Oxalobacteraceae, identical or closely related to strains with antifungal activity. Shoot and root communities significantly differed from each other but shared major OTUs. Watering but not warming restructured the seedling endophyte communities. Conclusions: The results suggest differences in assembly and ecological function across conifer life stages. Seedlings may recruit endophytes to protect against fungi under increased soil moisture.

Published 2017. This article is a U.S. Government work and is in the public domain in the USA Predictions of upslope range shifts for tree species with warming are based on assumptions of moisture stress at lower elevation limits and low-temperature stress at high-elevation limits. However, recent studies have shown that warming can reduce tree seedling establishment across the entire gradient from subalpine forest to alpine via moisture limitation. Warming effects also vary with species, potentially resulting in community shifts in high-elevation forests. We examined the growth and physiology underlying effects of warming on seedling demographic patterns. We evaluated dry mass (DM), root length, allocation above- and below-ground, and relative growth rate (RGR) of whole seedlings, and their ability to avoid or endure water stress via water-use efficiency and resisting turgor loss, for Pinus flexilis, Picea engelmannii and Pinus contorta seeded below, at and above tree line in experimentally warmed, watered and control plots in the Rocky Mountains, USA. We expected that growth and allocation responses to warming would relate to moisture status and that variation in drought tolerance traits would explain species differences in survival rates. Across treatments and elevations, seedlings of all species had weak turgor-loss resistance, and growth was marginal with negative RGR in the first growth phase (−0.01 to −0.04 g g−1 day−1). Growth was correlated with soil moisture, particularly in the relatively small-seeded P. contorta and P. engelmannii. Pinus flexilis, known to have the highest survivorship, attained the greatest DM and longest root but was also the slowest growing and most water-use efficient. This was likely due to its greater reliance on seed reserves. Seedlings developed 15% less total DM, 25% less root DM and 11% shorter roots in heated compared with unheated plots. Higher temperatures slightly increased DM, root length and RGR where soils were wettest, but more strongly decreased these variables under drier conditions. Synthesis. The surprising heat inhibition of tree seedling establishment at the cold edge of forests appears to have a physiological basis: newly germinated seedlings have poor moisture stress tolerance, which appears related to marginal initial growth and heavy reliance on seed reserves. Variation in these attributes among tree species at tree line helps explain their different climate responses.

© 2017 John Wiley & Sons Ltd Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here, we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had more than three-fold greater recruitment to their third year than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating halved recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering more than doubled recruitment, alleviating some of the negative effects of heating. Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid postfire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species’ range could enhance recruitment and facilitate upslope range shifts with climate change.

© 2017 Elsevier B.V. Metadata describe the ancillary information needed for data preservation and independent interpretation, comparison across heterogeneous datasets, and quality assessment and quality control (QA/QC). Environmental observations are vastly diverse in type and structure, can be taken across a wide range of spatiotemporal scales in a variety of measurement settings and approaches, and saved in multiple formats. Thus, well-organized, consistent metadata are required to produce usable data products from diverse environmental observations collected across field sites. However, existing metadata reporting protocols do not support the complex data synthesis and model-data integration needs of interdisciplinary earth system research. We developed a metadata reporting framework (FRAMES) to enable management and synthesis of observational data that are essential in advancing a predictive understanding of earth systems. FRAMES utilizes best practices for data and metadata organization enabling consistent data reporting and compatibility with a variety of standardized data protocols. We used an iterative scientist-centered design process to develop FRAMES, resulting in a data reporting format that incorporates existing field practices to maximize data-entry efficiency. Thus, FRAMES has a modular organization that streamlines metadata reporting and can be expanded to incorporate additional data types. With FRAMES's multi-scale measurement position hierarchy, data can be reported at observed spatial resolutions and then easily aggregated and linked across measurement types to support model-data integration. FRAMES is in early use by both data originators (persons generating data) and consumers (persons using data and metadata). In this paper, we describe FRAMES, identify lessons learned, and discuss areas of future development.

© 2017 by the Ecological Society of America. Mountain meadows have high biodiversity and help regulate stream water release following the snowmelt pulse. However, many meadows are experiencing woody plant encroachment, threatening these ecosystem services. While there have been field surveys of individual meadows and remote sensing-based landscape-scale studies of encroachment, what is missing is a broad-scale, ground-based study to understand common regional drivers, especially at high elevations, where land management has often played a less direct role. With this study, we ask: What are the climate and landscape conditions conducive to woody plant encroachment at the landscape scale, and how has historical climate variation affected tree recruitment in subalpine meadows over time? We measured density of encroaching trees across 340 subalpine meadows in the central Sierra Nevada, California, USA, and used generalized additive models (GAMs) to determine the relationship between landscape-scale patterns of encroachment and meadow environmental properties. We determined ages of trees in 30 survey meadows, used observed climate and GAMs to model the relationship between timing of recruitment and climate since the early 1900s, and extrapolated recruitment patterns into the future using downscaled climate scenarios. Encroachment was high among meadows with lodgepole pine (Pinus contorta Douglas ex Loudon var. murrayana (Balf.) Engelm.) in the immediate vicinity, at lower elevations, with physical conditions favoring strong soil drying, and with maximum temperatures above or below average. Climatic conditions during the year of germination were unimportant, with tree recruitment instead depending on a 3-yr seed production period prior to germination and a 6-yr seedling establishment period following germination. Recruitment was high when the seed production period had high snowpack, and when the seedling establishment period had warm summer maximum temperatures, high summer precipitation, and high snowpack. Applying our temporal model to downscaled output from four global climate models indicated that the average meadow will shift to forest by the end of the 21st century. Sierra Nevada meadow encroachment by conifers is ubiquitous and associated with climate conditions increasingly favorable for tree recruitment, which will lead to substantial changes in subalpine meadows and the ecosystem services they provide.

Species distribution shifts in response to climate change require that recruitment increase beyond current range boundaries. For trees with long life spans, the importance of climate‐sensitive seedling establishment to the pace of range shifts has not been demonstrated quantitatively. Using spatially explicit, stochastic population models combined with data from long‐term forest surveys, we explored whether the climate‐sensitivity of recruitment observed in climate manipulation experiments was sufficient to alter populations and elevation ranges of two widely distributed, high‐elevation North American conifers. Empirically observed, warming‐driven declines in recruitment led to rapid modelled population declines at the low‐elevation, ‘warm edge’ of subalpine forest and slow emergence of populations beyond the high‐elevation, ‘cool edge’. Because population declines in the forest occurred much faster than population emergence in the alpine, we observed range contraction for both species. For Engelmann spruce, this contraction was permanent over the modelled time horizon, even in the presence of increased moisture. For limber pine, lower sensitivity to warming may facilitate persistence at low elevations – especially in the presence of increased moisture – and rapid establishment above tree line, and, ultimately, expansion into the alpine. Synthesis. Assuming 21st century warming and no additional moisture, population dynamics in high‐elevation forests led to transient range contractions for limber pine and potentially permanent range contractions for Engelmann spruce. Thus, limitations to seedling recruitment with warming can constrain the pace of subalpine tree range shifts.

© 2016 by the Ecological Society of America. Climate change is expected to alter primary production and community composition in alpine ecosystems, but the direction and magnitude of change is debated. Warmer, wetter growing seasons may increase productivity; however, in the absence of additional precipitation, increased temperatures may decrease soil moisture, thereby diminishing any positive effect of warming. Since plant species show individual responses to environmental change, responses may depend on community composition and vary across life form or functional groups. We warmed an alpine plant community at Niwot Ridge, Colorado continuously for four years to test whether warming increases or decreases productivity of life form groups and the whole community. We provided supplemental water to a subset of plots to alleviate the drying effect of warming. We measured annual above-ground productivity and soil temperature and moisture, from which we calculated soil degree days and adequate soil moisture days. Using an information-theoretic approach, we observed that positive productivity responses to warming at the community level occur only when warming is combined with supplemental watering; otherwise we observed decreased productivity. Watering also increased community productivity in the absence of warming. Forbs accounted for the majority of the productivity at the site and drove the contingent community response to warming, while cushions drove the generally positive response to watering and graminoids muted the community response. Warming advanced snowmelt and increased soil degree days, while watering increased adequate soil moisture days. Heated and watered plots had more adequate soil moisture days than heated plots. Overall, measured changes in soil temperature and moisture in response to treatments were consistent with expected productivity responses. We found that available soil moisture largely determines the responses of this forb-dominated alpine community to simulated climate warming.

Coniferous forest nitrogen (N) budgets indicate unknown sources of N. A consistent association between limber pine (Pinus flexilis) and potential N2 -fixing acetic acid bacteria (AAB) indicates that native foliar endophytes may supply subalpine forests with N. To assess whether the P.flexilis-AAB association is consistent across years, we re-sampled P.flexilis twigs at Niwot Ridge, CO and characterized needle endophyte communities via 16S rRNA Illumina sequencing. To investigate whether endophytes have access to foliar N2 , we incubated twigs with (13) N2 -enriched air and imaged radioisotope distribution in needles, the first experiment of its kind using (13) N. We used the acetylene reduction assay to test for nitrogenase activity within P.flexilis twigs four times from June to September. We found evidence for N2 fixation in P.flexilis foliage. N2 diffused readily into needles and nitrogenase activity was positive across sampling dates. We estimate that this association could provide 6.8-13.6μgNm(-2) d(-1) to P.flexilis stands. AAB dominated the P.flexilis needle endophyte community. We propose that foliar endophytes represent a low-cost, evolutionarily stableN2 -fixing strategy for long-lived conifers. This novel source of biological N2 fixation has fundamental implications for understanding forest N budgets.

© 2015 New Phytologist Trust. Climate change is altering plant species distributions globally, and warming is expected to promote uphill shifts in mountain trees. However, at many cold-edge range limits, such as alpine treelines in the western United States, tree establishment may be colimited by low temperature and low moisture, making recruitment patterns with warming difficult to predict. We measured response functions linking carbon (C) assimilation and temperature- and moisture-related microclimatic factors for limber pine (Pinus flexilis) seedlings growing in a heating × watering experiment within and above the alpine treeline. We then extrapolated these response functions using observed microclimate conditions to estimate the net effects of warming and associated soil drying on C assimilation across an entire growing season. Moisture and temperature limitations were each estimated to reduce potential growing season C gain from a theoretical upper limit by 15-30% (c. 50% combined). Warming above current treeline conditions provided relatively little benefit to modeled net assimilation, whereas assimilation was sensitive to either wetter or drier conditions. Summer precipitation may be at least as important as temperature in constraining C gain by establishing subalpine trees at and above current alpine treelines as seasonally dry subalpine and alpine ecosystems continue to warm.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in under-represented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.