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  1. Abstract

    Resilience is the ability of ecosystems to maintain function while experiencing perturbation. Globally, forests are experiencing disturbances of unprecedented quantity, type, and magnitude that may diminish resilience. Early warning signals are statistical properties of data whose increase over time may provide insights into decreasing resilience, but there have been few applications to forests. We quantified four early warning signals (standard deviation, lag-1 autocorrelation, skewness, and kurtosis) across detrended time series of multiple ecosystem state variables at the Hubbard Brook Experimental Forest, New Hampshire, USA and analyzed how these signals have changed over time. Variables were collected over periods from 25 to 55 years from both experimentally manipulated and reference areas and were aggregated to annual timesteps for analysis. Long-term (>50 year) increases in early warning signals of stream calcium, a key biogeochemical variable at the site, illustrated declining resilience after decades of acid deposition, but only in watersheds that had previously been harvested. Trends in early warning signals of stream nitrate, a critical nutrient and water pollutant, likewise exhibited symptoms of declining resilience but in all watersheds. Temporal trends in early warning signals of some of groups of trees, insects, and birds also indicated changing resilience, but this pattern differed among, and even within, groups. Overall, ∼60% of early warning signals analyzed indicated decreasing resilience. Most of these signals occurred in skewness and kurtosis, suggesting ‘flickering’ behavior that aligns with emerging evidence of the forest transitioning into an oligotrophic condition. The other ∼40% of early warning signals indicated increasing or unchanging resilience. Interpretation of early warning signals in the context of system specific knowledge is therefore essential. They can be useful indicators for some key ecosystem variables; however, uncertainties in other variables highlight the need for further development of these tools in well-studied, long-term research sites.

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  2. Upslope shifts in plant distributions are often attributed to warming climate and lengthening of the growing season; however, biotic interactions may also contribute. The impacts of pests and pathogens are often sensitive to climate change and can vary along the climatic gradient associated with elevation. American beech ( Fagus grandifolia) has moved upslope throughout the northeastern United States. Meanwhile, beech growth and longevity have decreased as a result of beech bark disease (BBD), a decline disease caused by the introduced European felted beech scale insect ( Cryptococcus fagisuga) and native fungi from the genus Neonectria. Within a forested landscape spanning 250–1150 m elevation, we examined the relationships between elevation, beech demography and BBD to explore whether release from BBD at higher elevation may contribute to the upslope expansion of beech. Beech has shifted upslope at a rate of 1 m⋅year −1 coincident with lower mortality, higher recruitment, faster growth, lower BBD severity, and higher sapling densities at higher elevations. We suggest that climatic constraints on the beech scale insect at high elevations has led to a lower impact of BBD, which contributed to higher rates of beech growth, survival, and recruitment and in turn facilitated the regional upslope shift of beech. 
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  3. The understory layer is complex and includes groups of stems with distinctly different chances of survival and recruitment to the sapling size class. We explored how calcium amendment has impacted the trajectory of the seedling bank at Hubbard Brook Experimental Forest. The density of all tree stems in the seedling bank in 2018 (19 years after treatment) was greater in CAL (Watershed 1; calcium treatment) than REF (Waterhsed 6; reference) and beech was more abundant than sugar maple in both watersheds. In terms of relative abundance, the treatment had the opposite effects on the two species: the relative density of sugar maple was significantly greater in CAL than REF while the relative density of beech was significantly less. In terms of beech stem origin, Beech sprouts were more abundant than seedlings on both watersheds; however, beech stems of seed origin were more abundant on CAL (mean±1SE: 4.06±0.49 seedlings m-2) than REF (2.98±0.42), while sprouts were fewer (CAL: 14.4±1.30; REF: 20.5±1.47) resulting in the seedling to sprout ratio on CAL (1:3.5) being half that on REF (1:7). The influence on the seedling bank on future composition of these forests remains to be seen. These data were gathered as part of the Hubbard Brook Ecosystem Study (HBES). The HBES is a collaborative effort at the Hubbard Brook Experimental Forest, which is operated and maintained by the USDA Forest Service, Northern Research Station. 
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  4. The forest inventory surveys in the bird area were initiated in 1981 and transects were made permanent in 1991 by Tom Siccama who created and designed this tree survey. The inventory is representative of approximately 2.5 km2 of mid elevation northern hardwood forest. The data set is particularly geared toward producing accurate mortality and recruitment estimates. It consists of a total inventory of all trees greater than or equal to 10 cm dbh within each of four 10 m wide belt transects. The parallel transects are placed approximately 200 m apart and 290° bearing in an east-west direction for 2200 to 2900 m. In 1991, each live stem greater than or equal to 10 cm dbh was tagged with a unique number. Tree vigor is assessed every two years and diameter is remeasured every ten years. Every two years, new tags are placed on stems that have grown into the 10 cm diameter class. A survey of smaller trees (greater than or equal to 2 to less than 10 cm dbh) was first taken in 1991 and is resurveyed every ten years. This dataset includes 1991 and subsequent samplings. Data from an earlier sampling in 1981 can be found in: Sherry, T., D. Holmes, and T. Siccama. 2019. Forest Inventory of a Northern Hardwood Forest: Bird Area at the Hubbard Brook Experimental Forest, 1981 ver 7. Environmental Data Initiative. (Accessed 2021-09-16). These data were gathered as part of the Hubbard Brook Ecosystem Study (HBES). The HBES is a collaborative effort at the Hubbard Brook Experimental Forest, which is operated and maintained by the USDA Forest Service, Northern Research Station. These data have been used in the following publication: Siccama, T.G., Fahey, T.J., Johnson, C.E., Sherry, T.W., Denny, E.G., Girdler, E.B., Likens, G.E., and Schwarz, P.A. 2007. Population and biomass dynamics of trees in a northern hardwood forest at Hubbard Brook. Can. J. For. Res. 37(4): 737–749. doi:10.1139/X06-261. 
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  5. The valley-wide plots are a grid of 431 sites along fifteen N–S transects established at 500-m intervals spanning the entire Hubbard Brook Valley. The plot network was designed by Paul Schwarz for spatial analysis of tree species distribution patterns within the valley. Multiple above- and below-ground attributes have been measured on these plots. This dataset includes forest inventory data at 10 year intervals, for 1995, 2005, and 2015. The full survey takes three seasons to complete, with the datatable listing the exact measurement interval for each tree. Data are included for both trees and saplings on 371 core plots (all surveys) and 60 densified plots (1998, 2008). Locations of plots in this study can be found in the following dataset: Hubbard Brook Experimental Forest Valleywide Plots: GIS Shapefile (2022.) These data were gathered as part of the Hubbard Brook Ecosystem Study (HBES). The HBES is a collaborative effort at the Hubbard Brook Experimental Forest, which is operated and maintained by the USDA Forest Service, Northern Research Station. These data have been used in a number of publications including: Schwarz, P.A., Fahey, T.J., Martin, C.W., Siccama, T.G., and Bailey, A. 2001. Structure and composition of three northern hardwood–conifer forests with differing disturbance histories. For. Ecol. Manage. 144(1–3): 201–212. doi:10.1016/S0378-1127(00)00371-6. Schwarz, P.A., Fahey, T.J., and McCulloch, C.E. 2003. Factors controlling spatial variation of tree species abundance in a forested landscape. Ecology, 84(7): 1862–1878. doi:10.1890/0012-9658(2003)084[1862:FCSVOT]2.0.CO;2. van Doorn, N.S., Battles, J.J., Fahey, T.J., Siccama, T.G., and Schwarz, P.A. 2011. Links between biomass and tree demography in a northern hardwood forest: a decade of stability and change in Hubbard Brook Valley, New Hampshire. Can. J. For. Res. 41(7): 1369–1379. doi:10.1139/X11-063. Cleavitt, NL; AB Clyne and TJ Fahey. 2019. Epiphytic macrolichen patterns along an elevation gradient in the White Mountain National Forest, New Hampshire. J. Torrey Bot. Soc. 146(1): 8-17. Cleavitt, NL; Battles, JJ, Fahey, TJ, and Blum, J. 2014. Determinants of survival over seven years for a natural cohort of sugar maple seedlings in a northern hardwood forest. Can. J. For. Res.44 (9): 1112-1121. 
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  6. Abstract

    Range shifts of infectious plant disease are expected under climate change. As plant diseases move, emergent abiotic-biotic interactions are predicted to modify their distributions, leading to unexpected changes in disease risk. Evidence of these complex range shifts due to climate change, however, remains largely speculative. Here, we combine a long-term study of the infectious tree disease, white pine blister rust, with a six-year field assessment of drought-disease interactions in the southern Sierra Nevada. We find that climate change between 1996 and 2016 moved the climate optimum of the disease into higher elevations. The nonlinear climate change-disease relationship contributed to an estimated 5.5 (4.4–6.6) percentage points (p.p.) decline in disease prevalence in arid regions and an estimated 6.8 (5.8–7.9) p.p. increase in colder regions. Though climate change likely expanded the suitable area for blister rust by 777.9 (1.0–1392.9) km2into previously inhospitable regions, the combination of host-pathogen and drought-disease interactions contributed to a substantial decrease (32.79%) in mean disease prevalence between surveys. Specifically, declining alternate host abundance suppressed infection probabilities at high elevations, even as climatic conditions became more suitable. Further, drought-disease interactions varied in strength and direction across an aridity gradient—likely decreasing infection risk at low elevations while simultaneously increasing infection risk at high elevations. These results highlight the critical role of aridity in modifying host-pathogen-drought interactions. Variation in aridity across topographic gradients can strongly mediate plant disease range shifts in response to climate change.

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  8. null (Ed.)
    Accurate estimation of forest biomass is important for scientists and policymakers interested in carbon accounting, nutrient cycling, and forest resilience. Estimates often rely on the allometry of trees; however, limited datasets, uncertainty in model form, and unaccounted for sources of variation warrant a re-examination of allometric relationships using modern statistical techniques. We asked the following questions: (1) Is there among-stand variation in allometric relationships? (2) Is there nonlinearity in allometric relationships? (3) Can among-stand variation or nonlinearities in allometric equations be attributed to differences in stand age? (4) What are the implications for biomass estimation? To answer these questions, we synthesized a dataset of small trees from six different studies in the White Mountains of New Hampshire. We compared the performance of generalized additive models (GAMs) and linear models and found that GAMs consistently outperform linear models. The best-fitting model indicates that allometries vary among both stands and species and contain subtle nonlinearities which are themselves variable by species. Using a planned contrasts analysis, we were able to attribute some of the observed among-stand heterogeneity to differences in stand age. However, variability in these results point to additional sources of stand-level heterogeneity, which if identified could improve the accuracy of live-tree biomass estimation. 
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