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1. Root‐associated fungi not tree density influences stand nitrogen dynamics at the larch forest–tundra ecotone

   https://doi.org/10.1111/1365-2745.13882  

   Hewitt, Rebecca E. ; Alexander, Heather D. ; Miller, Samantha N. ; Mack, Michelle C. ( April 2022 , Journal of Ecology)

   Greater tree density and forest productivity at the tundra–taiga ecotone (TTE) are expected with climate warming, with potential feedbacks to the climate system. Yet, competition for nitrogen (N) may impact TTE dynamics. Greater tree density will likely increase N demand, while reducing N supply through soil shading and slower decomposition rates. We explored whether characteristics of roots and root‐associated fungi important to N acquisition responded to changes in density at the TTE and were related to above‐ground stand productivity and N cycling.

   We measured rooting depth, uptake of N forms among soil layers and ectomycorrhizal (EcM) colonization and composition along a natural tree density gradient of monodominant larchLarix cajanderiin northeastern Siberia. We tested relationships between larch root and fungal characteristics, above‐ground productivity and stand‐level N cycling parameters.

   Overall, there was preferential uptake of ammonium compared to glycine or nitrate. Nitrogen uptake was greatest in shallow soils of the organic horizon and related to root chemistry, root‐associated fungi and above‐ground N cycling parameters, but the direction of these relationships depended on N form. Uptake of different N forms, rooting depth and EcM colonization and composition were not related to tree density, but fungal composition was correlated with root N chemistry and above‐ground N cycling parameters. In addition to EcM, the abundance of dark septate endophytes and other ascomycetous taxa was positively related to N uptake and above‐ground N cycling parameters.

   Synthesis. There was little impact of tree density on root and fungal parameters related to N acquisition suggesting intraspecific larch competition for N was not amplified with increased density. There was, however, a strong impact of root‐associated fungi on N uptake and stand N dynamics regardless of tree density. Together, this suggests an important role of root‐associated fungi on broadscale patterns of N cycling in TTE larch forests independent of changes in tree density expected with climate warming.

    

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2. Increasing tree density accelerates stand‐level nitrogen cycling at the taiga–tundra ecotone in northeastern Siberia

   https://doi.org/10.1002/ecs2.4175  

   Hewitt, Rebecca E. ; Alexander, Heather D. ; Izbicki, Brian ; Loranty, Michael M. ; Natali, Susan M. ; Walker, Xanthe J. ; Mack, Michelle C. ( July 2022 , Ecosphere)

   As climate warms, tree density at the taiga–tundra ecotone (TTE) is expected to increase, which may intensify competition for belowground resources in this nitrogen (N)‐limited environment. To determine the impacts of increased tree density on N cycling and productivity, we examined edaphic properties indicative of soil N availability along with aboveground and belowground tree‐level traits and stand characteristics related to carbon (C) and N cycling across a tree density gradient of monodominant larch (Larix cajanderi) at the TTE in far northeastern Siberia. We found no consistent evidence from soil, tree, or stand‐level N cycling characteristics of lower N availability or greater intraspecific competition for N with increased density. Active layer thickness declined, but resin‐sorbed N and soil organic layer thickness did not covary with increased tree density. There was, however, greater allocation belowground to stand‐level coarse and fine roots with increased tree density, an allocation pattern suggestive of limited soil resources. Foliar traits related to C (%C, δ¹³C, and resorption) were responsive to density indicating the importance of non‐nutrient resources, like light, to foliar stoichiometry. As tree density increased and individual trees had lower productivity, tree‐level N and biomass pools aboveground and belowground declined tracking decreases in N uptake, N resorption, N use efficiency, and allocation to slow cycling tissues like wood. At the stand level, our findings show high N turnover with increased N acquisition, allocation to short‐lived tissues with relatively high N content and reduced N residence time, and greater stand productivity as tree density increased. Yet, these positive relationships were curtailed at the highest tree densities. Our observations of shifts in biomass, C and N allocation, and loss aboveground, along with greater root density with increased tree density, could have strong impacts on C and N cycling and should be represented in models of TTE dynamics and feedbacks to climate.

    

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3. Tree density influences ecohydrological drivers of plant–water relations in a larch boreal forest in Siberia

   https://doi.org/10.1002/eco.2132  

   Kropp, Heather ; Loranty, Michael M. ; Natali, Susan M. ; Kholodov, Alexander L. ; Alexander, Heather D. ; Zimov, Nikita S. ; Mack, Michelle C. ; Spawn, Seth A. ( July 2019 , Ecohydrology)

   Transpiration and stomatal conductance in deciduous needleleaf boreal forests of northern Siberia can be highly sensitive to water stress, permafrost thaw, and atmospheric dryness. Additionally, north‐eastern Siberian boreal forests are fire‐driven, and larch (Larixspp.) are the sole tree species. We examined differences in tree water use, stand characteristics, and stomatal responses to environmental drivers between high and low tree density stands that burned 76 years ago in north‐eastern Siberia. Our results provide process‐level insight to climate feedbacks related to boreal forest productivity, water cycles, and permafrost across Arctic regions. The high density stand had shallower permafrost thaw depths and deeper moss layers than the low density stand. Rooting depths and shallow root biomass were similar between stands. Daily transpiration was higher on average in the high‐density stand 0.12 L m⁻² day⁻¹(SE: 0.004) compared with the low density stand 0.10 L m⁻² day⁻¹(SE: 0.001) throughout the abnormally wet summer of 2016. Transpiration rates tended to be similar at both stands during the dry period in 2017 in both stands of 0.10 L m⁻² day⁻¹(SE: 0.002). The timing of precipitation impacted stomatal responses to environmental drivers, and the high density stand was more dependent on antecedent precipitation that occurred over longer periods in the past compared with the low density stand. Post‐fire tree density differences in plant–water relations may lead to different trajectories in plant mortality, water stress, and ecosystem water cycles across Siberian landscapes.

    

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4. Increasing rates of subalpine tree mortality linked to warmer and drier summers

   https://doi.org/10.1111/1365-2745.13634  

   Andrus, Robert A. ; Chai, Rachel K. ; Harvey, Brian J. ; Rodman, Kyle C. ; Veblen, Thomas T. ; Battipaglia, ed., Giovanna ( March 2021 , Journal of Ecology)

   Warming temperatures and rising moisture deficits are expected to increase the rates of background tree mortality–low amounts of tree mortality (~0.5%–2% year⁻¹), characterizing the forest demographic processes in the absence of abrupt, coarse‐scale disturbance events (e.g. fire). When compounded over multiple decades and large areas, even minor increases in background tree mortality (e.g. <0.5% year⁻¹) can cause changes to forest communities and carbon storage potential that are comparable to or greater than those caused by disturbances.

   We examine how temporal variability in rates of background tree mortality for four subalpine conifers reflects variability in climate and climate teleconnections using observations of tree mortality from 1982 to 2019 at Niwot Ridge, Colorado, USA. Individually marked trees (initial population 5,043) in 13 permanent plots—located across a range of site conditions, stand ages and species compositions—were censused for new mortality nine times over 37 years.

   Background tree mortality was primarily attributed to stress from unfavourable climate and competition (71.2%) and bark beetle activity (23.3%), whereas few trees died from wind (5.3%) and wildlife impacts (0.2%). Mean annualized tree mortality attributed to tree stress and bark beetles more than tripled across all stands between initial censuses (0.26% year⁻¹, 1982–1993/1994) and recent censuses (0.82% year⁻¹, 2008–2019). Higher rates of tree mortality were related to warmer maximum summer temperatures, greater summer moisture deficits, and negative anomalies in ENSO (La Niña), with greater effects of drought in some subpopulations (tree size, age and species). For example, in older stands (>250 years), larger and older trees were more likely to die than smaller and younger trees. Differences in tree mortality rates and sensitivity to climate among subpopulations that varied by stand type may lead to unexpected shifts in stand composition and structure.

   Synthesis. A strong relationship between higher rates of tree mortality and warmer, drier summer climate conditions implies that climate warming will continue to increase background mortality rates in subalpine forests. Combined with increases in disturbances and declining frequency of moist‐cool years suitable for seedling establishment, increasing rates of tree mortality have the potential to drive declines in subalpine tree populations.

    

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5. Understory plant diversity and composition across a postfire tree density gradient in a Siberian Arctic boreal forest

   https://doi.org/10.1139/cjfr-2020-0483  

   Paulson, Alison K. ; Peña, Homero ; Alexander, Heather D. ; Davydov, Sergei P. ; Loranty, Michael M. ; Mack, Michelle C. ; Natali, Susan M. ( May 2021 , Canadian Journal of Forest Research)

   null (Ed.)

   Cajander larch (Larix cajanderi Mayr.) forests of the Siberian Arctic are experiencing increased wildfire activity in conjunction with climate warming. These shifts could affect postfire variation in the density and arrangement of trees and understory plant communities. To better understand how understory plant composition, abundance, and diversity vary with tree density, we surveyed understory plant communities and stand characteristics (e.g., canopy cover, active layer depth, and soil organic layer depth) within 25 stands representing a density gradient of similarly-aged larch trees that established following a 1940 fire near Cherskiy, Russia. Understory plant diversity and mean total plant abundance decreased with increased canopy cover. Canopy cover was also the most important variable affecting individual species’ abundances. In general, tall shrubs (e.g., Betula nana subsp. exilis) were more abundant in low-density stands with high light availability, and mosses (e.g., Sanionia spp.) were more abundant in high-density stands with low light availability. These results provide evidence that postfire variation in tree recruitment affects understory plant community composition and diversity as stands mature. Therefore, projected increases in wildfire activity in the Siberian Arctic could have cascading impacts on forest structure and composition in both overstory and understory plant communities. 

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