Arctic sea ice retreat fuels boreal forest advance.

Climate-induced northward advance of boreal forest is expected to lessen albedo, alter carbon stocks, and replace tundra, but where and when this advance will occur remains largely unknown. Using data from 19 sites across 22 degrees of longitude along the tree line of northern Alaska, we show a stronger temporal correlation of tree ring growth with open water uncovered by retreating Arctic sea ice than with air temperature. Spatially, our results suggest that tree growth, recruitment, and range expansion are causally linked to open water through associated warmer temperatures, deeper snowpacks, and improved nutrient availability. We apply a meta-analysis to 82 circumarctic sites, finding that proportionally more tree lines have advanced where proximal to ongoing sea ice loss. Taken together, these findings underpin how and where changing sea ice conditions facilitate high-latitude forest advance.

Root-associated fungi and acquisitive root traits facilitate permafrost nitrogen uptake from long-term experimentally warmed tundra.

Root-associated fungi (RAF) and root traits regulate plant acquisition of nitrogen (N), which is limiting to growth in Arctic ecosystems. With anthropogenic warming, a new N source from thawing permafrost has the potential to change vegetation composition and increase productivity, influencing climate feedbacks. Yet, the impact of warming on tundra plant root traits, RAF, and access to permafrost N is uncertain. We investigated the relationships between RAF, species-specific root traits, and uptake of N from the permafrost boundary by tundra plants experimentally warmed for nearly three decades at Toolik Lake, Alaska. Warming increased acquisitive root traits of nonmycorrhizal and mycorrhizal plants. RAF community composition of ericoid (ERM) but not ectomycorrhizal (ECM) shrubs was impacted by warming and correlated with root traits. RAF taxa in the dark septate endophyte, ERM, and ECM guilds strongly correlated with permafrost N uptake for ECM and ERM shrubs. Overall, a greater proportion of variation in permafrost N uptake was related to root traits than RAF. Our findings suggest that warming Arctic ecosystems will result in interactions between roots, RAF, and newly thawed permafrost that may strongly impact feedbacks to the climate system through mechanisms of carbon and N cycling.

Sufficient conditions for rapid range expansion of a boreal conifer.

Unprecedented modern rates of warming are expected to advance boreal forest into Arctic tundra1, thereby reducing albedo2-4, altering carbon cycling4 and further changing climate1-4, yet the patterns and processes of this biome shift remain unclear5. Climate warming, required for previous boreal advances6-17, is not sufficient by itself for modern range expansion of conifers forming forest-tundra ecotones5,12-15,17-20. No high-latitude population of conifers, the dominant North American Arctic treeline taxon, has previously been documented5 advancing at rates following the last glacial maximum (LGM)6-8. Here we describe a population of white spruce (Picea glauca) advancing at post-LGM rates7 across an Arctic basin distant from established treelines and provide evidence of mechanisms sustaining the advance. The population doubles each decade, with exponential radial growth in the main stems of individual trees correlating positively with July air temperature. Lateral branches in adults and terminal leaders in large juveniles grow almost twice as fast as those at established treelines. We conclude that surpassing temperature thresholds1,6-17, together with winter winds facilitating long-distance dispersal, deeper snowpack and increased soil nutrient availability promoting recruitment and growth, provides sufficient conditions for boreal forest advance. These observations enable forecast modelling with important insights into the environmental conditions converting tundra into forest.

Microbiome assembly in thawing permafrost and its feedbacks to climate.

The physical and chemical changes that accompany permafrost thaw directly influence the microbial communities that mediate the decomposition of formerly frozen organic matter, leading to uncertainty in permafrost-climate feedbacks. Although changes to microbial metabolism and community structure are documented following thaw, the generality of post-thaw assembly patterns across permafrost soils of the world remains uncertain, limiting our ability to predict biogeochemistry and microbial community responses to climate change. Based on our review of the Arctic microbiome, permafrost microbiology, and community ecology, we propose that Assembly Theory provides a framework to better understand thaw-mediated microbiome changes and the implications for community function and climate feedbacks. This framework posits that the prevalence of deterministic or stochastic processes indicates whether the community is well-suited to thrive in changing environmental conditions. We predict that on a short timescale and following high-disturbance thaw (e.g., thermokarst), stochasticity dominates post-thaw microbiome assembly, suggesting that functional predictions will be aided by detailed information about the microbiome. At a longer timescale and lower-intensity disturbance (e.g., active layer deepening), deterministic processes likely dominate, making environmental parameters sufficient for predicting function. We propose that the contribution of stochastic and deterministic processes to post-thaw microbiome assembly depends on the characteristics of the thaw disturbance, as well as characteristics of the microbial community, such as the ecological and phylogenetic breadth of functional guilds, their functional redundancy, and biotic interactions. These propagate across space and time, potentially providing a means for predicting the microbial forcing of greenhouse gas feedbacks to global climate change.

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.

Mycobiont contribution to tundra plant acquisition of permafrost-derived nitrogen.

As Arctic soils warm, thawed permafrost releases nitrogen (N) that could stimulate plant productivity and thus offset soil carbon losses from tundra ecosystems. Although mycorrhizal fungi could facilitate plant access to permafrost-derived N, their exploration capacity beyond host plant root systems into deep, cold active layer soils adjacent to the permafrost table is unknown. We characterized root-associated fungi (RAF) that colonized ericoid (ERM) and ectomycorrhizal (ECM) shrub roots and occurred below the maximum rooting depth in permafrost thaw-front soil in tussock and shrub tundra communities. We explored the relationships between root and thaw front fungal composition and plant uptake of a 15 N tracer applied at the permafrost boundary. We show that ERM and ECM shrubs associate with RAF at the thaw front providing evidence for potential mycelial connectivity between roots and the permafrost boundary. Among shrubs and tundra communities, RAF connectivity to the thaw boundary was ubiquitous. The occurrence of particular RAF in both roots and thaw front soil was positively correlated with 15 N recovered in shrub biomass Taxon-specific RAF associations could be a mechanism for the vertical redistribution of deep, permafrost-derived nutrients, which may alleviate N limitation and stimulate productivity in warming tundra.

Poor nutrition as a potential cause of divergent tree growth near the Arctic treeline in northern Alaska.

Trees growing near the Arctic treeline have long been used to reconstruct past climates. However, recent studies have shown deterioration of historically strong positive correlations between air temperature and tree growth (known as "divergence"). Divergence has important implications for confidence in paleoclimate reconstructions and ecosystem-atmosphere carbon exchange. Studies in the Brooks Range of northern Alaska showed that white spruce in the west increased growth in response to late 20th century warming, whereas those in the east failed to show a growth increase. In an earlier study across four watersheds in the Brooks Range, we tested and rejected the hypothesis that divergence in the easternmost watershed reflects moisture limitation of growth. Here, using 16 sites distributed across the same four watersheds, we tested an alternative hypothesis, that greater nutrient limitation in the east may have impeded positive growth responses to warming. Climate comparison across the four Brooks Range study watersheds revealed that, although the easternmost watershed generally had a drier growing-season climate, the most consistent difference was that winter air temperature and both winter and summer soil temperatures were much colder in the central and eastern watersheds. Soil nutrient availability, foliar nutrient concentrations, and tree growth were all generally lower in the central and eastern than in the western watersheds. Foliar phosphorus concentration was the best predictor of spatial variation in branch extension growth-a finding that is somewhat inconsistent with the theory that forest productivity on young, glacially derived soils should be strongly nitrogen limited. Experimental fertilization yielded the greatest growth increase in the eastern, an intermediate response in the central, and the smallest growth increase in the western watershed, generally mirroring trends in soil temperature, soil nutrient availability, foliar nutrient concentrations, and growth of control trees. Our results confirm that growth in the easternmost watershed is more nutrient limited and suggest that phosphorus limitation may be at least as important as nitrogen limitation of growth. We hypothesize that cold soil effects on tree access to nutrients might explain divergence in the eastern Brooks Range and elsewhere near the Arctic treeline, particularly in areas with cold winters and widespread permafrost.

Variable retention harvesting influences belowground plant-fungal interactions of Nothofagus pumilio seedlings in forests of southern Patagonia.

BACKGROUND: The post-harvest recovery and sustained productivity of Nothofagus pumilio forests in Tierra del Fuego may be affected by the abundance and composition of ectomycorrhizal fungi (EMF). Timber harvesting alters EMF community structure in many managed forests, but the impacts of harvesting can vary with the management strategy. The implementation of variable retention (VR) management can maintain, increase, or decrease the diversity of many species, but the effects of VR on EMF in the forests of southern Patagonia have not been studied, nor has the role of EMF in the regeneration process of these forests. METHODS: We evaluated the effects of VR management on the EMF community associated with N. pumilio seedlings. We quantified the abundance, composition, and diversity of EMF across aggregate (AR) and dispersed (DR) retention sites within VR managed areas, and compared them to primary forest (PF) unmanaged stands. EMF assemblage and taxonomic identities were determined by ITS-rDNA sequencing of individual root tips sampled from 280 seedlings across three landscape replicates. To better understand seedling performance, we tested the relationships between EMF colonization, EMF taxonomic composition, seedling biomass, and VR treatment. RESULTS: The majority of EMF taxa were Basidiomycota belonging to the families Cortinariaceae (n = 29), Inocybaceae (n = 16), and Thelephoraceae (n = 8), which was in agreement with other studies of EMF diversity in Nothofagus forests. EMF richness and colonization was reduced in DR compared to AR and PF. Furthermore, EMF community composition was similar between AR and PF, but differed from the composition in DR. EMF community composition was correlated with seedling biomass and soil moisture. The presence of Peziza depressa was associated with higher seedling biomass and greater soil moisture, while Inocybe fibrillosibrunnea and Cortinarius amoenus were associated with reduced seedling biomass and lower soil moisture. Seedling biomass was more strongly related to retention type than EMF colonization, richness, or composition. DISCUSSION: Our results demonstrate reduced EMF attributes and altered composition in VR treatments relative to PF stands, with stronger impacts in DR compared to AR. This suggests that VR has the potential to improve the conservation status of managed stands by supporting native EMF in AR. Our results also demonstrate the complex linkages between retention treatments, fungal community composition, and tree growth at individual and stand scales.

The potential for mycobiont sharing between shrubs and seedlings to facilitate tree establishment after wildfire at Alaska arctic treeline.

Root-associated fungi, particularly ectomycorrhizal fungi (EMF), are critical symbionts of all boreal tree species. Although climatically driven increases in wildfire frequency and extent have been hypothesized to increase vegetation transitions from tundra to boreal forest, fire reduces mycorrhizal inoculum. Therefore, changes in mycobiont inoculum may potentially limit tree-seedling establishment beyond current treeline. We investigated whether ectomycorrhizal shrubs that resprout after fire support similar fungal taxa to those that associate with tree seedlings that establish naturally after fire. We then assessed whether mycobiont identity correlates with the biomass or nutrient status of these tree seedlings. The majority of fungal taxa observed on shrub and seedling root systems were EMF, with some dark septate endophytes and ericoid mycorrhizal taxa. Seedlings and adjacent shrubs associated with similar arrays of fungal taxa, and there were strong correlations between the structure of seedling and shrub fungal communities. These results show that resprouting postfire shrubs support fungal taxa compatible with tree seedlings that establish after wildfire. Shrub taxon, distance to the nearest shrub and fire severity influenced the similarity between seedling and shrub fungal communities. Fungal composition was correlated with both foliar C:N ratio and seedling biomass and was one of the strongest explanatory variables predicting seedling biomass. While correlative, these results suggest that mycobionts are important to nutrient acquisition and biomass accrual of naturally establishing tree seedlings at treeline and that mycobiont taxa shared by resprouting postfire vegetation may be a significant source of inoculum for tree-seedling establishment beyond current treeline.

Fire-severity effects on plant-fungal interactions after a novel tundra wildfire disturbance: implications for arctic shrub and tree migration.

BACKGROUND: Vegetation change in high latitude tundra ecosystems is expected to accelerate due to increased wildfire activity. High-severity fires increase the availability of mineral soil seedbeds, which facilitates recruitment, yet fire also alters soil microbial composition, which could significantly impact seedling establishment. RESULTS: We investigated the effects of fire severity on soil biota and associated effects on plant performance for two plant species predicted to expand into Arctic tundra. We inoculated seedlings in a growth chamber experiment with soils collected from the largest tundra fire recorded in the Arctic and used molecular tools to characterize root-associated fungal communities. Seedling biomass was significantly related to the composition of fungal inoculum. Biomass decreased as fire severity increased and the proportion of pathogenic fungi increased. CONCLUSIONS: Our results suggest that effects of fire severity on soil biota reduces seedling performance and thus we hypothesize that in certain ecological contexts fire-severity effects on plant-fungal interactions may dampen the expected increases in tree and shrub establishment after tundra fire.