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1. 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)

   Abstract 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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2. Positive response of tree productivity to warming is reversed by increased tree density at the Arctic tundra-taiga ecotone

   https://doi.org/10.1139/cjfr-2020-0466  

   Walker, Xanthe; Alexander, Heather D.; Berner, Logan; Boyd, Melissa A; Loranty, Michael M.; Natali, Sue; Mack, Michelle C. (January 2021, Canadian Journal of Forest Research)

   null (Ed.)

   The transition zone between the northern boreal forest and the arctic tundra, known as the tundra-taiga ecotone (TTE) has undergone rapid warming in recent decades. In response to this warming, tree density, growth, and stand productivity are expected to increase. Increases in tree density have the potential to negate the positive impacts of warming on tree growth through a reduction in the active layer and an increase in competitive interactions. We assessed the effects of tree density on tree growth and climate-growth responses of Cajander larch (Larix cajanderi) and on trends in the normalized difference vegetation index (NDVI) in the TTE of Northeast Siberia. We examined 19 mature forest stands that all established after a fire in 1940 and ranged in tree density from 300 to 37,000 stems ha-1. High density stands with shallow active layers had lower tree growth, higher stand productivity, and more negative growth responses to growing season temperatures compared to low density stands with deep active layers. Variation in stand productivity across the density gradient was not captured by Landsat derived NDVI, but NDVI did capture annual variations in stand productivity. Our results suggest that the expected increases in tree density following fires at the TTE may effectively limit tree growth and that NDVI is unlikely to capture increasing productivity associated with changes in tree density. 

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3. 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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4. 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)

   Abstract 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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5. Linking Post-fire Tree Density to Carbon Storage in High-Latitude Cajander Larch (Larix cajanderi) Forests of Far Northeastern Siberia

   https://doi.org/10.1007/s10021-024-00913-0  

   Alexander, H_D; Paulson, A_K; Loranty, M_M; Mack, M_C; Natali, S_M; Pena, H.; Davydov, S.; Spektor, V.; Zimov, N. (May 2024, Ecosystems)

   Abstract With climate warming and drying, fire activity is increasing in Cajander larch (Larix cajanderiMayr.) forests underlain by continuous permafrost in northeastern Siberia, and initial post-fire tree demographic processes could unfold to determine long-term forest carbon (C) dynamics through impacts on tree density. Here, we evaluated above- and belowground C pools across 25 even-aged larch stands of varying tree densities that established following a wildfire in ~ 1940 near Cherskiy, Russia. Total C pools increased with increased larch tree density, from ~ 9,000 g C m⁻²in low-density stands to ~ 11,000 g C m⁻²in high and very high-density stands, with increases most pronounced at tree densities < 1 stem m⁻²and driven by increased above- and belowground (that is, coarse roots) and live and dead (that is, woody debris and snags) larch biomass. Total understory vegetation and non-larch coarse root C pools declined with increased tree density due to decreased shrub C pools, but these pools were relatively small compared to larch biomass. Fine root, soil organic matter (OM), and near surface (0–30 cm) mineral soil (MS) C pools varied little with tree density, although soil C pools held most (18–28% in OM and 44–51% in MS) C stored in these stands. Thus, if changing fire regimes promote denser stands, C storage will likely increase, but whether this increase offsets C lost during fires remains unknown. Our findings highlight how post-fire tree demographic processes impact C pool distribution and stability in larch forests of Siberian permafrost regions. 

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