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Resource allocation theory posits that increased soil nutrient availability results in decreased plant investment in nutrient acquisition. We evaluated this theory by quantifying fine root biomass and growth in a long term, nitrogen (N) × phosphorus (P) fertilization study in three mature northern hardwood forest stands where aboveground growth increased primarily in response to P addition. We did not detect a decline in fine root biomass or growth in response to either N or P. Instead, fine root growth increased in response to N, by 40% for length (P = 0.04 for the main effect of N in ANOVA), and by 36% for mass, relative to controls. Fine root mass growth was lower in response to N + P addition than predicted from the main effects of N and P (P = 0.01 for the interaction of N × P). The response of root growth to N availability did not result in detectable responses in fine root biomass (P = 0.61), which is consistent with increased root turnover with N addition. We propose that the differential growth response to fertilization between above- and belowground components is a mechanism by which trees enhance P acquisition in response to increasing N availability, illustrating how both elements may co-limit northern hardwood forest production.

Functional balance theory predicts that plants will allocate less carbon belowground when the availability of nutrients is elevated. We tested this prediction in two successional northern hardwood forest stands by quantifying fine root biomass and growth after 5–7 years of treatment in a nitrogen (N) x phosphorus (P) factorial addition experiment. We quantified root responses at two different levels of treatment: the whole-plot scale fertilization and small-patch scale fertilization of ingrowth cores. Fine root biomass was higher in plots receiving P, and fine root growth was highest in plots receiving both N and P. Thus, belowground productivity did not decrease in response to long-term addition of nutrients. We did not find conclusive evidence that elevated availability of one nutrient at the plot scale induced foraging for the other nutrient at the core scale, or that foraging for nutrients at the core scale responded to addition of limiting nutrients. Our observations suggest NP co-limitation of fine root growth and indicate complex interactions of N and P affecting aboveground and belowground production in early successional northern hardwood forest ecosystems.

ContextIncreased anthropogenic climate forcing is projected to have tremendous impacts on global forest ecosystems, with northern biomes being more at risk.ObjectivesTo model the impacts of harvest and increased anthropogenic climate forcing on eastern Canada’s forest landscapes and to assess the strong spatial heterogeneity in the severity, the nature and direction of the impacts expected within northern forest regions.MethodsWe used LANDIS-II to project species-specific aboveground biomass (AGB) between 2020 and 2150 under three climate (baseline, RCP 4.5 and RCP 8.5) and two harvest (baseline harvest, no harvest) scenarios within four forest regions (boreal west, boreal east, mixedwood and northern hardwood).ResultsClimate change impacts within the boreal forest regions would mainly result from increases in wildfires activity which will strongly alter total AGB. In the mixedwood and northern hardwood, changes will be less important and will result from climate-induced growth constraints that will alter species composition towards more thermophilous species. Climate-induced impacts were much more important and swifter under RCP 8.5 after 2080 suggesting that eastern Canada’s forests might cross important tipping points under strong anthropogenic climate forcing.ConclusionsBoreal forest regions will be much less resilient than mixedwood or northern hardwoods to the projected changes in climate regimes. Current harvest strategies will interact with anthropogenic climate forcing to further modify forest landscapes, notably by accelerating thermophilous species AGB gain in southernmost regions. Major changes to harvest practices are strongly needed to preserve the long-term sustainability of wood supply in eastern Canada. Adaptation strategies should be region-specific.

Soil microbes mediate major biogeochemical processes in forest ecosystems. Soil pH is considered a “master variable” with a strong positive effect on many biogeochemical processes. To better understand how soil pH influences microbial activity and nitrogen (N) dynamics in forests, we utilized a set of long-term measurements of surface soil pH, N availability, and microbial biomass and respiration from the Hubbard Brook Experimental Forest (HBEF), a northern hardwood forest in New Hampshire, USA. We compared the strengths of these relationships in an unmanipulated watershed, where naturally acidic soils have been further acidified by anthropogenic acid deposition, to those in a nearby watershed, where soils were treated with calcium silicate to ameliorate the effects of acid deposition. While we expected to observe strong positive relationships between soil pH and microbial biomass and activity, we instead found weak and/or curvilinear relationships. In many cases, microbial biomass and activity peaked at unexpectedly low pH values (~ 4.5), and decreased at higher pH values, especially in the calcium-treated soils. It is likely that complexities in plant-microbial interactions inhibit and/or mask microbial response to changes in pH in these acidic soils. These results raise questions about pH as a controller of microbial processes and how ecosystems recover in response to decreases in acid deposition.

Over the next century, many mid and high latitude temperate ecosystems are projected to experience rising growing season temperatures and increased frequency of soil freeze/thaw cycles (FTCs) due to a reduction in the depth and duration of the winter snowpack. We conducted a manipulative field experiment in a northern hardwood forest at the Hubbard Brook Experimental Forest in New Hampshire to determine the interactive effects of climate change across seasons on rates of net N mineralization, foliar N, and natural abundance foliar ¹⁵N (δ¹⁵N) in red maple (Acer rubrum) trees. We warmed soils 5 °C above ambient temperatures and induced winter FTCs to simulate projected changes over the next century. Net N mineralization was dominated by ammonification and increased with warmer soil temperatures, but was not affected by soil FTCs in the previous winter. Similarly, warming led to increased foliar N concentrations and δ¹⁵N, with no effect of soil FTCs. Together, our results show that growing season soil warming increases soil N availability and N uptake by trees, which may offset the previously observed negative effects of a smaller snowpack and more frequent soil freezing on N cycling. We conclude that soil warming in the growing season may counteract the trend of reduced soil N availability relative to plant N demand (i.e. N oligotrophication) observed in northern hardwood forests. This research demonstrates that climate change across seasons affects N cycling in northern hardwood forests in ways that would have not been apparent from examining one season alone.

Tick-borne pathogens pose a significant risk to livestock, wildlife and public health. Host-seeking behaviours may depend on a combination of infection status and environmental factors. Here, we assessed the effects of habitat type and pathogen infection on host-seeking behaviour (questing) in the lone star tick, Amblyomma americanum. Ticks were collected using a tick drag from two different habitat types: xeric hammock and successional hardwood forests. Using a standardized assay, we recorded the likelihood of questing for each tick, the average height quested and total time spent questing and then tested each tick for the presence of Rickettsia spp. and Ehrlichia spp. using conventional polymerase chain reaction. We did not detect Ehrlichia in any ticks, although 30% tested positive for Rickettsia amblyommatis, a member of the Rickettsia spotted fever group. Ticks infected with R. amblyommatis spent less time questing compared to uninfected ticks, with infected ticks spending 85 s on average questing and uninfected ticks spending 112 s. Additionally, ticks collected from xeric hammock habitats spent over twice as long questing compared to ticks from successional hardwood forests. Ticks from xeric hammock spent 151 s on average questing while ticks from successional hardwood forest spent only 58 s during a 10-min observation period. These results demonstrate that habitat type and infection status can influence tick host-seeking behaviours, which can play a pivotal role in disease dynamics.

Detecting latitudinal range shifts of forest trees in response to recent climate change is difficult because of slow demographic rates and limited dispersal but may be facilitated by spatially compressed climatic zones along elevation gradients in montane environments. We resurveyed forest plots established in 1964 along elevation transects in the Green Mountains (Vermont) to examine whether a shift had occurred in the location of the northern hardwood-boreal forest ecotone (NBE) from 1964 to 2004. We found a 19% increase in dominance of northern hardwoods from 70% in 1964 to 89% in 2004 in the lower half of the NBE. This shift was driven by a decrease (up to 76%) in boreal and increase (up to 16%) in northern hardwood basal area within the lower portions of the ecotone. We used aerial photographs and satellite imagery to estimate a 91- to 119-m upslope shift in the upper limits of the NBE from 1962 to 2005. The upward shift is consistent with regional climatic change during the same period; interpolating climate data to the NBE showed a 1.1°C increase in annual temperature, which would predict a 208-m upslope movement of the ecotone, along with a 34% increase in precipitation. The rapid upward movement of the NBE indicates little inertia to climatically induced range shifts in montane forests; the upslope shift may have been accelerated by high turnover in canopy trees that provided opportunities for ingrowth of lower elevation species. Our results indicate that high-elevation forests may be jeopardized by climate change sooner than anticipated.

Wood is a hygroscopic material that can reach an equilibrium moisture content when ambient temperature and relative humidity are constant. Moisture affects all properties of wood, as well as its preservative treatment. The hygroscopic behavior of wood can be attributed to the hydroxyl groups of its constituents. Since hemicellulose shows the greatest water affinity, it can be considered the main responsible for the ingress of water into the wood mass. Below the fiber saturation point, wood moisture is only stored in the cell walls. Proton Nuclear Magnetic Resonance (NMR) is a relative method used for the evaluation of moisture content distribution in wood and NMR relaxation is an excellent tool to study the hygroscopic behavior of different woods below the fiber saturation point. This work aimed to test the hypothesis of discriminating among softwoods and hardwoods of different botanical species and identifying further sub-clusters of woods based on the NMR proton spin–spin ( T 2 ) and spin–lattice ( T 1 ) relaxation times of their cell wall water in the hygroscopic moisture range. Importantly, the study was performed using a portable low-cost NMR instrument with which it is possible to investigate wood samples of any size. The main result of this study was that at RH = 94% the relaxation time T 2,2 , associated with the cell wall bound water, can be used as a marker to discriminate among softwoods and hardwoods. Graphical abstract

Climate models project higher growing-season temperatures and a decline in the depth and duration of winter snowpack throughout many north temperate ecosystems over the next century. A smaller snowpack is projected to induce more frequent soil freeze/thaw cycles in winter in northern hardwood forests of the northeastern United States. We measured the combined effects of warmer growing-season soil temperatures and increased winter freeze/thaw cycles on rates of leaf-level photosynthesis and transpiration (sap flow) of red maple (Acer rubrum) trees in a northern hardwood forest at the Climate Change Across Seasons Experiment at Hubbard Brook Experimental Forest in New Hampshire. Soil temperatures were warmed 5°C above ambient temperatures during the growing season and soil freeze/thaw cycles were induced in winter to mimic the projected changes in soil temperature over the next century. Relative to reference plots, growing-season soil warming increased rates of leaf-level photosynthesis by up to 85.32 ± 4.33%, but these gains were completely offset by soil freeze/thaw cycles in winter, suggesting that increased freeze/thaw cycles in winter over the next 100 yr will reduce the effect of warming on leaf-level carbon gains. Soil warming in the growing season increased rates of transpiration per kilopascal of vapor pressure deficit (VPD) by up to 727.39 ± 0.28%, even when trees were exposed to increased frequency of soil freeze/thaw cycles in the previous winter, which could influence regional hydrology in the future. Using climate projections downscaled from the Coupled Model Intercomparison Project, we project increased rates of whole-season transpiration in these forests over the next century by 42–61%. We also project 52–77 additional days when daily air temperatures will be above the long-term average daily maximum during the growing season at Hubbard Brook. Together, these results show that projected changes in climate across both the growing season and winter are likely to cause greater rates of water uptake and have no effect on rates of leaf-level carbon uptake by trees, with potential ecosystem consequences for hydrology and carbon cycling in northern hardwood forests.