Insights into the tussock growth form with model-data fusion.

Some rhizomatous grass and sedge species form tussocks that impact ecosystem structure and function. Despite their importance, tussock development and size controls are poorly understood due to the decadal to centennial timescales over which tussocks form. We explored mechanisms regulating tussock development and size in a ubiquitous arctic tussock sedge (Eriophorum vaginatum) using field observations and a mass balance model coupled with a tiller population model. Model-data fusion was used to quantify parameter and prediction uncertainty, determine model sensitivity, and test hypotheses on the factors regulating tussock size. The model accurately captured the dynamics of tussock development, characteristics, and size observed in the field. Tussock growth approached maximal size within several decades, which was determined by feedbacks between the mass balance of tussock root necromass and density-dependent tillering. The model also predicted that maximal tussock size was primarily regulated by tiller root productivity and necromass bulk density and less so by tiller demography. These predictions were corroborated by field observations of tussock biomass and root characteristics. The study highlights the importance of belowground processes in regulating tussock development and size and enhances our understanding of the influence of tussocks on arctic ecosystem structure and function.

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.

Cross-biome synthesis of source versus sink limits to tree growth.

Uncertainties surrounding tree carbon allocation to growth are a major limitation to projections of forest carbon sequestration and response to climate change. The prevalence and extent to which carbon assimilation (source) or cambial activity (sink) mediate wood production are fundamentally important and remain elusive. We quantified source-sink relations across biomes by combining eddy-covariance gross primary production with extensive on-site and regional tree ring observations. We found widespread temporal decoupling between carbon assimilation and tree growth, underpinned by contrasting climatic sensitivities of these two processes. Substantial differences in assimilation-growth decoupling between angiosperms and gymnosperms were determined, as well as stronger decoupling with canopy closure, aridity, and decreasing temperatures. Our results reveal pervasive sink control over tree growth that is likely to be increasingly prominent under global climate change.

Small herbivores with big impacts: Tundra voles (Microtus oeconomus) alter post-fire ecosystem dynamics.

Fire is an important ecological disturbance that can reset ecosystems and initiate changes in plant community composition, ecosystem biogeochemistry, and primary productivity. As herbivores rely on primary producers for food, changes in vegetation may alter plant-herbivore interactions with important-but often unexplored-feedbacks to ecosystems. Here we examined the impact of post-fire changes in plant community composition and structure on habitat suitability and rodent herbivore activity in response to a large, severe, and unprecedented fire in northern Alaskan tundra. In moist acidic tundra where the fire occurred, tundra voles (Microtus oeconomus) are the dominant herbivore and rely on the tussock forming sedge Eriophorum vaginatum for both food and nesting material. Tundra voles were 10 times more abundant at the burned site compared with nearby unburned tundra 7-12 years after the fire. Fire increased the habitat suitability for voles by increasing plant productivity and biomass, food quality, and cover through both taller vegetation and increased microtopography. As a result of elevated vole abundance, Eriophorum mortality caused by vole herbivory was two orders of magnitude higher than natural mortality and approached the magnitude of the mortality rate resulting directly from the fire. These findings suggest that post-fire increases in herbivore pressure on Eriophorum could, in turn, disrupt graminoid recovery and enhance shrub encroachment. Tundra state transitions from graminoid to shrub dominated are also evident following other disturbances and fertilization experiments, suggesting that as Arctic temperatures rise, greater available nutrients and increased frequencies of large-scale disturbances may also alter plant-animal interactions with cascading impacts on plant communities and ecosystem function.

Alleviation of nutrient co-limitation induces regime shifts in post-fire community composition and productivity in Arctic tundra.

Recent unprecedented fires in the Arctic during the past two decades have indicated a pressing need to understand the long-term ecological impacts of fire in this biome. Anecdotal evidence suggests that tundra fires can induce regime shifts that change tussock tundra to more shrub-dominated ecosystems. However, the ecological mechanisms regulating these shifts are poorly understood, but are hypothesized to involve changes to nutrient availability in this nutrient limited system. Here we conducted a 4-year two-factorial (control: C, nitrogen along: N+ , phosphorus alone: P+ , nitrogen and phosphorus combined: NP+ ) fertilization experiment in both unburned and burned tundra to test this hypothesis after a decade of post-fire recovery. A decade after fire, the burned site exhibited an increase in soil nitrogen and phosphorus availability and a transition toward taller, more productive, and more deciduous vegetation. This shift in vegetation structure, composition, and function was induced at the unburned site through the addition of both NP+ and the alleviation of their co-limitation. Both burned and unburned tundra responded similarly to fertilizer treatments by increasing leaf area index, greenness, and canopy height in NP+ treatments, and exhibited no significant response in individual N+ or P+ treatments. These results point to a greater need to understand coupled carbon, nitrogen, and phosphorus cycles in this system, and suggest that post-fire regime shifts are regulated by the alleviation of nitrogen and phosphorus co-limitation in Arctic tundra.

Tundra wildfire triggers sustained lateral nutrient loss in Alaskan Arctic.

Climate change is creating widespread ecosystem disturbance across the permafrost zone, including a rapid increase in the extent and severity of tundra wildfire. The expansion of this previously rare disturbance has unknown consequences for lateral nutrient flux from terrestrial to aquatic environments. Lateral loss of nutrients could reduce carbon uptake and slow recovery of already nutrient-limited tundra ecosystems. To investigate the effects of tundra wildfire on lateral nutrient export, we analyzed water chemistry in and around the 10-year-old  Anaktuvuk River fire scar in northern Alaska. We collected water samples from 21 burned and 21 unburned watersheds during snowmelt, at peak growing season, and after plant senescence in 2017 and 2018. After a decade of ecosystem recovery, aboveground biomass had recovered in burned watersheds, but overall carbon and nitrogen remained ~20% lower, and the active layer remained ~10% deeper. Despite lower organic matter stocks, dissolved organic nutrients were substantially elevated in burned watersheds, with higher flow-weighted concentrations of organic carbon (25% higher), organic nitrogen (59% higher), organic phosphorus (65% higher), and organic sulfur (47% higher). Geochemical proxies indicated greater interaction with mineral soils in watersheds with surface subsidence, but optical analysis and isotopes suggested that recent plant growth, not mineral soil, was the main source of organic nutrients in burned watersheds. Burned and unburned watersheds had similar δ15 N-NO3 - , indicating that exported nitrogen was of preburn origin (i.e., not recently fixed). Lateral nitrogen flux from burned watersheds was 2- to 10-fold higher than rates of background nitrogen fixation and atmospheric deposition estimated in this area. These findings indicate that wildfire in Arctic tundra can destabilize nitrogen, phosphorus, and sulfur previously stored in permafrost via plant uptake and leaching. This plant-mediated nutrient loss could exacerbate terrestrial nutrient limitation after disturbance or serve as an important nutrient release mechanism during succession.

An Open-Source, Durable, and Low-Cost Alternative to Commercially Available Soil Temperature Data Loggers.

Soil temperatures play an important role in determining the distribution and function of organisms. However, soil temperature is decoupled from air temperature and varies widely in space. Characterizing and predicting soil temperature requires large and expensive networks of data loggers. We developed an open-source soil temperature data logger and created online resources to ensure our design was accessible. We tested data loggers constructed by students, with little prior electronics experience, in the lab, and in the field in Alaska. The do-it-yourself (DIY) data logger was comparably accurate to a commercial system with a mean absolute error of 2% from -20-0 °C and 1% from 0-20 °C. They captured accurate soil temperature data and performed reliably in the field with less than 10% failing in the first year of deployment. The DIY loggers were ~1.7-7 times less expensive than commercial systems. This work has the potential to increase the spatial resolution of soil temperature monitoring and serve as a powerful educational tool. The DIY soil temperature data logger will reduce data collection costs and improve our understanding of species distributions and ecological processes. It also provides an educational resource to enhance STEM, accessibility, inclusivity, and engagement.

Limited overall impacts of ectomycorrhizal inoculation on recruitment of boreal trees into Arctic tundra following wildfire belie species-specific responses.

We tested whether post-fire seedling establishment of common boreal tree and expanding shrub species at treeline and in Arctic tundra is facilitated by co-migration of boreal forest mycorrhizal fungi. Wildfires are anticipated to facilitate biome shifts at the forest-tundra ecotone by improving seedbed conditions for recruiting boreal species; at the same time fire alters the composition and availability of mycorrhizal fungi critical to seedling performance. To determine the role of root-associated fungi (RAF) in post-fire seedling recruitment and future biome shifts, we outplanted four dominant boreal tree and shrub species inoculated with one of three treatments at treeline and in tundra: burned boreal forest, unburned boreal forest, or a control treatment of sterilized inoculum. We compared survivorship, growth, and physiological performance of the seedlings in relation to mycorrhizal inoculum treatment and among host species, characterized the RAF communities based on ITS-rDNA sequencing of individual root tips sampled from surviving seedlings, and tested for correlations between RAF composition and the inoculation treatments, host species, and duration of the experiment. We explored correlations between RAF composition and seedling metrics. Both live and sterile autoclaved inoculation treatments had similar effects on seedling survivorship and growth for all species. RAF composition did not vary by treatment, suggesting that most colonization was due to local fungi. However, seedling traits and growth were correlated with RAF species composition, colonization, and the relative abundance of specific RAF taxa. Picea sp. performance in particular showed strong co-variation with RAF metrics. Our results suggest that mycorrhizal co-migration is not a primary limiting factor to boreal seedling recruitment because the experimental provision of inoculum did not affect seedling recruitment; yet, RAF did influence seedling performance, particularly resident RAF at treeline and in tundra, suggesting that mycorrhizal fungi are important to vegetation processes at the treeline-tundra ecotone.

Differential responses of ecotypes to climate in a ubiquitous Arctic sedge: implications for future ecosystem C cycling.

The response of vegetation to climate change has implications for the carbon cycle and global climate. It is frequently assumed that a species responds uniformly across its range to climate change. However, ecotypes - locally adapted populations within a species - display differences in traits that may affect their gross primary productivity (GPP) and response to climate change. To determine if ecotypes are important for understanding the response of ecosystem productivity to climate we measured and modeled growing season GPP in reciprocally transplanted and experimentally warmed ecotypes of the abundant Arctic sedge Eriophorum vaginatum. Transplanted northern ecotypes displayed home-site advantage in GPP that was associated with differences in leaf area index. Southern ecotypes exhibited a greater response in GPP when transplanted. The results demonstrate that ecotypic differentiation can impact the morphology and function of vegetation with implications for carbon cycling. Moreover they suggest that ecotypic control of GPP may limit the response of ecosystem productivity to climate change. This investigation shows that ecotypes play a substantial role in determining GPP and its response to climate. These results have implications for understanding annual to decadal carbon cycling where ecotypes could influence ecosystem function and vegetation feedbacks to climate change.

Solar-induced chlorophyll fluorescence exhibits a universal relationship with gross primary productivity across a wide variety of biomes.

In our recent study in Global Change Biology (Li et al., ), we examined the relationship between solar-induced chlorophyll fluorescence (SIF) measured from the Orbiting Carbon Observatory-2 (OCO-2) and gross primary productivity (GPP) derived from eddy covariance flux towers across the globe, and we discovered that there is a nearly universal relationship between SIF and GPP across a wide variety of biomes. This finding reveals the tremendous potential of SIF for accurately mapping terrestrial photosynthesis globally.

Solar-induced chlorophyll fluorescence is strongly correlated with terrestrial photosynthesis for a wide variety of biomes: First global analysis based on OCO-2 and flux tower observations.

Solar-induced chlorophyll fluorescence (SIF) has been increasingly used as a proxy for terrestrial gross primary productivity (GPP). Previous work mainly evaluated the relationship between satellite-observed SIF and gridded GPP products both based on coarse spatial resolutions. Finer resolution SIF (1.3 km × 2.25 km) measured from the Orbiting Carbon Observatory-2 (OCO-2) provides the first opportunity to examine the SIF-GPP relationship at the ecosystem scale using flux tower GPP data. However, it remains unclear how strong the relationship is for each biome and whether a robust, universal relationship exists across a variety of biomes. Here we conducted the first global analysis of the relationship between OCO-2 SIF and tower GPP for a total of 64 flux sites across the globe encompassing eight major biomes. OCO-2 SIF showed strong correlations with tower GPP at both midday and daily timescales, with the strongest relationship observed for daily SIF at the 757 nm (R2  = 0.72, p < 0.0001). Strong linear relationships between SIF and GPP were consistently found for all biomes (R2  = 0.57-0.79, p < 0.0001) except evergreen broadleaf forests (R2  = 0.16, p < 0.05) at the daily timescale. A higher slope was found for C4 grasslands and croplands than for C3 ecosystems. The generally consistent slope of the relationship among biomes suggests a nearly universal rather than biome-specific SIF-GPP relationship, and this finding is an important distinction and simplification compared to previous results. SIF was mainly driven by absorbed photosynthetically active radiation and was also influenced by environmental stresses (temperature and water stresses) that determine photosynthetic light use efficiency. OCO-2 SIF generally had a better performance for predicting GPP than satellite-derived vegetation indices and a light use efficiency model. The universal SIF-GPP relationship can potentially lead to more accurate GPP estimates regionally or globally. Our findings revealed the remarkable ability of finer resolution SIF observations from OCO-2 and other new or future missions (e.g., TROPOMI, FLEX) for estimating terrestrial photosynthesis across a wide variety of biomes and identified their potential and limitations for ecosystem functioning and carbon cycle studies.

Modeling long-term changes in tundra carbon balance following wildfire, climate change, and potential nutrient addition.

To investigate the underlying mechanisms that control long-term recovery of tundra carbon (C) and nutrients after fire, we employed the Multiple Element Limitation (MEL) model to simulate 200-yr post-fire changes in the biogeochemistry of three sites along a burn severity gradient in response to increases in air temperature, CO2 concentration, nitrogen (N) deposition, and phosphorus (P) weathering rates. The simulations were conducted for severely burned, moderately burned, and unburned arctic tundra. Our simulations indicated that recovery of C balance after fire was mainly determined by the internal redistribution of nutrients among ecosystem components (controlled by air temperature), rather than the supply of nutrients from external sources (e.g., nitrogen deposition and fixation, phosphorus weathering). Increases in air temperature and atmospheric CO2 concentration resulted in (1) a net transfer of nutrient from soil organic matter to vegetation and (2) higher C : nutrient ratios in vegetation and soil organic matter. These changes led to gains in vegetation biomass C but net losses in soil organic C stocks. Under a warming climate, nutrients lost in wildfire were difficult to recover because the warming-induced acceleration in nutrient cycles caused further net nutrient loss from the system through leaching. In both burned and unburned tundra, the warming-caused acceleration in nutrient cycles and increases in ecosystem C stocks were eventually constrained by increases in soil C : nutrient ratios, which increased microbial retention of plant-available nutrients in the soil. Accelerated nutrient turnover, loss of C, and increasing soil temperatures will likely result in vegetation changes, which further regulate the long-term biogeochemical succession. Our analysis should help in the assessment of tundra C budgets and of the recovery of biogeochemical function following fire, which is in turn necessary for the maintenance of wildlife habitat and tundra vegetation.

Modeling carbon-nutrient interactions during the early recovery of tundra after fire.

Fire frequency has dramatically increased in the tundra of northern Alaska, USA, which has major implications for the carbon budget of the region and the functioning of these ecosystems, which support important wildlife species. We investigated the postfire succession of plant and soil carbon (C), nitrogen (N), and phosphorus (P) fluxes and stocks along a burn severity gradient in the 2007 Anaktuvuk River fire scar in northern Alaska. Modeling results indicated that the early regrowth of postfire tundra vegetation was limited primarily by its canopy photosynthetic potential, rather than nutrient availability, because of the initially low leaf area and relatively high inorganic N and P concentrations in soil. Our simulations indicated that the postfire recovery of tundra vegetation was sustained predominantly by the uptake of residual inorganic N (i.e., in the remaining ash), and the redistribution of N and P from soil organic matter to vegetation. Although residual nutrients in ash were higher in the severe burn than the moderate burn, the moderate burn recovered faster because of the higher remaining biomass and consequent photosynthetic potential. Residual nutrients in ash allowed both burn sites to recover and exceed the unburned site in both aboveground biomass and production five years after the fire. The investigation of interactions among postfire C, N, and P cycles has contributed to a mechanistic understanding of the response of tundra ecosystems to fire disturbance. Our study provided insight on how the trajectory of recovery of tundra from wildfire is regulated during early succession.

Disturbance legacies and climate jointly drive tree growth and mortality in an intensively studied boreal forest.

Most North American forests are at some stage of post-disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short-term studies in a longer term context, and particularly important for the carbon-dense, fire-prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine-learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999-2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr-(1), with most mortality occurring in medium-sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current- and previous-year data exerted significant effects. Models based on these variables explained 23-44% of the ring-width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old-growth stands in a disturbance-prone landscape, and the sustainable management of regional forests in a changing climate.

The application of δ¹8O and δD for understanding water pools and fluxes in a Typha marsh.

The δ¹8O and δD composition of water pools (leaf, root, standing water and soil water) and fluxes [transpiration (T), evaporation (E)] were used to understand ecohydrological processes in a managed Typha latifolia L. freshwater marsh. We observed isotopic steady-state T and deep rooting in Typha. The isotopic mass balance of marsh standing water showed that E accounted for 3% of the total water loss, T accounted for 17% and subsurface drainage (D) accounted for the majority (80%). There was a vertical gradient in water vapour content and isotopic composition within and above the canopy sufficient for constructing an isotopic mass balance of water vapour during some sampling periods. During these periods, the proportion of T in evapotranspiration (T/ET) was between 56 ± 17% and 96 ± 67%, and the estimated error was relatively high (>37%) because of non-local, background sources in vapour. Independent estimates of T/ET using eddy covariance measurements yielded similar mean values during the Typha growing season. The various T/ET estimates agreed that T was the dominant source of marsh vapour loss in the growing season. The isotopic mass balance of water vapour yielded reasonable results, but the mass balance of standing water provided more definitive estimates of water losses.

Burn severity influences postfire CO2 exchange in arctic tundra.

Burned landscapes present several challenges to quantifying landscape carbon balance. Fire scars are composed of a mosaic of patches that differ in burn severity, which may influence postfire carbon budgets through damage to vegetation and carbon stocks. We deployed three eddy covariance towers along a burn severity gradient (i.e., severely burned, moderately burned, and unburned tundra) to monitor postfire net ecosystem exchange of CO2 (NEE) within the large 2007 Anaktuvuk River fire scar in Alaska, USA, during the summer of 2008. Remote sensing data from the MODerate resolution Imaging Spectroradiometer (MODIS) was used to assess the spatial representativeness of the tower sites and parameterize a NEE model that was used to scale tower measurements to the landscape. The tower sites had similar vegetation and reflectance properties prior to the Anaktuvuk River fire and represented the range of surface conditions observed within the fire scar during the 2008 summer. Burn severity influenced a variety of surface properties, including residual organic matter, plant mortality, and vegetation recovery, which in turn determined postfire NEE. Carbon sequestration decreased with increased burn severity and was largely controlled by decreases in canopy photosynthesis. The MODIS two-band enhanced vegetation index (EVI2) monitored the seasonal course of surface greenness and explained 86% of the variability in NEE across the burn severity gradient. We demonstrate that understanding the relationship between burn severity, surface reflectance, and NEE is critical for estimating the overall postfire carbon balance of the Anaktuvuk River fire scar.