The three major axes of terrestrial ecosystem function.

The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8.

Mechanistic evidence for tracking the seasonality of photosynthesis with solar-induced fluorescence.

Northern hemisphere evergreen forests assimilate a significant fraction of global atmospheric CO2 but monitoring large-scale changes in gross primary production (GPP) in these systems is challenging. Recent advances in remote sensing allow the detection of solar-induced chlorophyll fluorescence (SIF) emission from vegetation, which has been empirically linked to GPP at large spatial scales. This is particularly important in evergreen forests, where traditional remote-sensing techniques and terrestrial biosphere models fail to reproduce the seasonality of GPP. Here, we examined the mechanistic relationship between SIF retrieved from a canopy spectrometer system and GPP at a winter-dormant conifer forest, which has little seasonal variation in canopy structure, needle chlorophyll content, and absorbed light. Both SIF and GPP track each other in a consistent, dynamic fashion in response to environmental conditions. SIF and GPP are well correlated (R2 = 0.62-0.92) with an invariant slope over hourly to weekly timescales. Large seasonal variations in SIF yield capture changes in photoprotective pigments and photosystem II operating efficiency associated with winter acclimation, highlighting its unique ability to precisely track the seasonality of photosynthesis. Our results underscore the potential of new satellite-based SIF products (TROPOMI, OCO-2) as proxies for the timing and magnitude of GPP in evergreen forests at an unprecedented spatiotemporal resolution.

Seasonal variation in the canopy color of temperate evergreen conifer forests.

Evergreen conifer forests are the most prevalent land cover type in North America. Seasonal changes in the color of evergreen forest canopies have been documented with near-surface remote sensing, but the physiological mechanisms underlying these changes, and the implications for photosynthetic uptake, have not been fully elucidated. Here, we integrate on-the-ground phenological observations, leaf-level physiological measurements, near surface hyperspectral remote sensing and digital camera imagery, tower-based CO2 flux measurements, and a predictive model to simulate seasonal canopy color dynamics. We show that seasonal changes in canopy color occur independently of new leaf production, but track changes in chlorophyll fluorescence, the photochemical reflectance index, and leaf pigmentation. We demonstrate that at winter-dormant sites, seasonal changes in canopy color can be used to predict the onset of canopy-level photosynthesis in spring, and its cessation in autumn. Finally, we parameterize a simple temperature-based model to predict the seasonal cycle of canopy greenness, and we show that the model successfully simulates interannual variation in the timing of changes in canopy color. These results provide mechanistic insight into the factors driving seasonal changes in evergreen canopy color and provide opportunities to monitor and model seasonal variation in photosynthetic activity using color-based vegetation indices.

Montane forest productivity across a semiarid climatic gradient.

High-elevation montane forests are disproportionately important to carbon sequestration in semiarid climates where low elevations are dry and characterized by low carbon density ecosystems. However, these ecosystems are increasingly threatened by climate change with seasonal implications for photosynthesis and forest growth. As a result, we leveraged eddy covariance data from six evergreen conifer forest sites in the semiarid western United States to extrapolate the status of carbon sequestration within a framework of projected warming and drying. At colder locations, the seasonal evolution of gross primary productivity (GPP) was characterized by a single broad maximum during the summer that corresponded to snow melt-derived moisture and a transition from winter dormancy to spring activity. Conversely, winter dormancy was transient at warmer locations, and GPP was responsive to both winter and summer precipitation such that two distinct GPP maxima were separated by a period of foresummer drought. This resulted in a predictable sequence of primary limiting factors to GPP beginning with air temperature in winter and proceeding to moisture and leaf area during the summer. Due to counteracting winter (positive) and summer (negative) GPP responses to warming, leaf area index and moisture availability were the best predictors of annual GPP differences across sites. Overall, mean annual GPP was greatest at the warmest site due to persistent vegetation photosynthetic activity throughout the winter. These results indicate that the trajectory of this region's carbon sequestration will be sensitive to reduced or delayed summer precipitation, especially if coupled to snow drought and earlier soil moisture recession, but summer precipitation changes remain highly uncertain. Given the demonstrated potential for seasonally offsetting responses to warming, we project that decadal semiarid montane forest carbon sequestration will remain relatively stable in the absence of severe disturbance.

Ecosystem transpiration and evaporation: Insights from three water flux partitioning methods across FLUXNET sites.

We apply and compare three widely applicable methods for estimating ecosystem transpiration (T) from eddy covariance (EC) data across 251 FLUXNET sites globally. All three methods are based on the coupled water and carbon relationship, but they differ in assumptions and parameterizations. Intercomparison of the three daily T estimates shows high correlation among methods (R between .89 and .94), but a spread in magnitudes of T/ET (evapotranspiration) from 45% to 77%. When compared at six sites with concurrent EC and sap flow measurements, all three EC-based T estimates show higher correlation to sap flow-based T than EC-based ET. The partitioning methods show expected tendencies of T/ET increasing with dryness (vapor pressure deficit and days since rain) and with leaf area index (LAI). Analysis of 140 sites with high-quality estimates for at least two continuous years shows that T/ET variability was 1.6 times higher across sites than across years. Spatial variability of T/ET was primarily driven by vegetation and soil characteristics (e.g., crop or grass designation, minimum annual LAI, soil coarse fragment volume) rather than climatic variables such as mean/standard deviation of temperature or precipitation. Overall, T and T/ET patterns are plausible and qualitatively consistent among the different water flux partitioning methods implying a significant advance made for estimating and understanding T globally, while the magnitudes remain uncertain. Our results represent the first extensive EC data-based estimates of ecosystem T permitting a data-driven perspective on the role of plants' water use for global water and carbon cycling in a changing climate.

Joint control of terrestrial gross primary productivity by plant phenology and physiology.

Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate-carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy-covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000-2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r(2) = 0.90) and GPP recovery after a fire disturbance in South Dakota (r(2) = 0.88). Additional analysis of the eddy-covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Lake Superior evaporation: A long-term eddy covariance dataset at Stannard Rock Lighthouse (2008-2022).

Robust, accurate, and direct measurements of evaporation and related energy fluxes on the Laurentian Great Lakes are necessary to understand the large historical range in water levels, regional climatology, lake hydrodynamics, and lake-effect snowfall, all of which inform water management. Despite the societal and scientific importance of this information, few long-term, full-year, in situ measurements exist due to logistical, financial, and safety-related challenges. We present 15 years (2008-2022) of eddy covariance data from Stannard Rock, a historic lighthouse on Lake Superior located 38 km southeast of Manitou Island and 72 km north of Marquette, Michigan. We provide information about the site and instrumentation, as well as data availability and processing. Analysis of this unique long-term dataset, available through the AmeriFlux network (US-GL1), will improve our ability to understand the drivers and patterns of large-lake surface energy fluxes and will advance predictions of evaporative regimes over Lake Superior.

Bark beetle impacts on forest evapotranspiration and its partitioning.

Insect outbreaks affect forest structure and function and represent a major category of forest disturbance globally. However, the resulting impacts on evapotranspiration (ET), and especially hydrological partitioning between the abiotic (evaporation) and biotic (transpiration) components of total ET, are not well constrained. As a result, we combined remote sensing, eddy covariance, and hydrological modeling approaches to determine the effects of bark beetle outbreak on ET and its partitioning at multiple scales throughout the Southern Rocky Mountain Ecoregion (SRME), USA. At the eddy covariance measurement scale, 85 % of the forest was affected by beetles, and water year ET as a fraction of precipitation (P) decreased by 30 % relative to a control site, with 31 % greater reductions in growing season transpiration relative to total ET. At the ecoregion scale, satellite remote sensing masked to areas of >80 % tree mortality showed corresponding ET/P reductions of 9-15 % that occurred 6-8 years post-disturbance, and indicated that the majority of the total reduction occurred during the growing season; the Variable Infiltration Capacity hydrological model showed an associated 9-18 % increase in the ecoregion runoff ratio. Long-term (16-18 year) ET and vegetation mortality datasets extend the length of previously published analyses and allowed for clear characterization of the forest recovery period. During that time, transpiration recovery outpaced total ET recovery, which was lagged in part due to persistently reduced winter sublimation, and there was associated evidence of increasing late summer vegetation moisture stress. Overall, comparison of three independent methods and two partitioning approaches demonstrated a net negative impact of bark beetles on ET, and a relatively greater negative impact on transpiration, following bark beetle outbreak in the SRME.

Vegetation type is an important predictor of the arctic summer land surface energy budget.

Despite the importance of high-latitude surface energy budgets (SEBs) for land-climate interactions in the rapidly changing Arctic, uncertainties in their prediction persist. Here, we harmonize SEB observations across a network of vegetated and glaciated sites at circumpolar scale (1994-2021). Our variance-partitioning analysis identifies vegetation type as an important predictor for SEB-components during Arctic summer (June-August), compared to other SEB-drivers including climate, latitude and permafrost characteristics. Differences among vegetation types can be of similar magnitude as between vegetation and glacier surfaces and are especially high for summer sensible and latent heat fluxes. The timing of SEB-flux summer-regimes (when daily mean values exceed 0 Wm-2) relative to snow-free and -onset dates varies substantially depending on vegetation type, implying vegetation controls on snow-cover and SEB-flux seasonality. Our results indicate complex shifts in surface energy fluxes with land-cover transitions and a lengthening summer season, and highlight the potential for improving future Earth system models via a refined representation of Arctic vegetation types.

Temperature thresholds of ecosystem respiration at a global scale.

Ecosystem respiration is a major component of the global terrestrial carbon cycle and is strongly influenced by temperature. The global extent of the temperature-ecosystem respiration relationship, however, has not been fully explored. Here, we test linear and threshold models of ecosystem respiration across 210 globally distributed eddy covariance sites over an extensive temperature range. We find thresholds to the global temperature-ecosystem respiration relationship at high and low air temperatures and mid soil temperatures, which represent transitions in the temperature dependence and sensitivity of ecosystem respiration. Annual ecosystem respiration rates show a markedly reduced temperature dependence and sensitivity compared to half-hourly rates, and a single mid-temperature threshold for both air and soil temperature. Our study indicates a distinction in the influence of environmental factors, including temperature, on ecosystem respiration between latitudinal and climate gradients at short (half-hourly) and long (annual) timescales. Such climatological differences in the temperature sensitivity of ecosystem respiration have important consequences for the terrestrial net carbon sink under ongoing climate change.

The FLUXNET2015 dataset and the ONEFlux processing pipeline for eddy covariance data.

The FLUXNET2015 dataset provides ecosystem-scale data on CO2, water, and energy exchange between the biosphere and the atmosphere, and other meteorological and biological measurements, from 212 sites around the globe (over 1500 site-years, up to and including year 2014). These sites, independently managed and operated, voluntarily contributed their data to create global datasets. Data were quality controlled and processed using uniform methods, to improve consistency and intercomparability across sites. The dataset is already being used in a number of applications, including ecophysiology studies, remote sensing studies, and development of ecosystem and Earth system models. FLUXNET2015 includes derived-data products, such as gap-filled time series, ecosystem respiration and photosynthetic uptake estimates, estimation of uncertainties, and metadata about the measurements, presented for the first time in this paper. In addition, 206 of these sites are for the first time distributed under a Creative Commons (CC-BY 4.0) license. This paper details this enhanced dataset and the processing methods, now made available as open-source codes, making the dataset more accessible, transparent, and reproducible.

Evidence for non-steady-state carbon emissions from snow-scoured alpine tundra.

High-latitude warming is capable of accelerating permafrost degradation and the decomposition of previously frozen carbon. The existence of an analogous high-altitude feedback, however, has yet to be directly evaluated. We address this knowledge gap by coupling a radiocarbon-based model to 7 years (2008-2014) of continuous eddy covariance data from a snow-scoured alpine tundra meadow in Colorado, USA, where solifluction lobes are associated with discontinuous permafrost. On average, the ecosystem was a net annual source of 232 ± 54 g C m-2 (mean ± 1 standard deviation) to the atmosphere, and respiration of relatively radiocarbon-depleted (i.e., older) substrate contributes to carbon emissions during the winter. Given that alpine soils with permafrost occupy 3.6 × 106 km2 land area and are estimated to contain 66.3 Pg of soil organic carbon (4.5% of the global pool), this scenario has global implications for the mountain carbon balance and corresponding resource allocation to lower elevations.

Short-term favorable weather conditions are an important control of interannual variability in carbon and water fluxes.

Ecosystem models often perform poorly in reproducing interannual variability in carbon and water fluxes, resulting in considerable uncertainty when estimating the land-carbon sink. While many aggregated variables (growing season length, seasonal precipitation, or temperature) have been suggested as predictors for interannual variability in carbon fluxes, their explanatory power is limited and uncertainties remain as to their relative contributions. Recent results show that the annual count of hours where evapotranspiration (ET) is larger than its 95th percentile is strongly correlated with the annual variability of ET and gross primary production (GPP) in an ecosystem model. This suggests that the occurrence of favorable conditions has a strong influence on the annual carbon budget. Here we analyzed data from eight forest sites of the AmeriFlux network with at least 7 years of continuous measurements. We show that for ET and the carbon fluxes GPP, ecosystem respiration (RE), and net ecosystem production, counting the "most active hours/days" (i.e., hours/days when the flux exceeds a high percentile) correlates well with the respective annual sums, with correlation coefficients generally larger than 0.8. Phenological transitions have much weaker explanatory power. By exploiting the relationship between most active hours and interannual variability, we classify hours as most active or less active and largely explain interannual variability in ecosystem fluxes, particularly for GPP and RE. Our results suggest that a better understanding and modeling of the occurrence of large values in high-frequency ecosystem fluxes will result in a better understanding of interannual variability of these fluxes.