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The goal of this project is to test the overarching hypothesis that positive feedback mechanisms involving changes in seasonal cycles that diminish N availability to plants such that plant N demand is not met by soil N availability in northern forests. Specifically, we hypothesize that increasing N demand by plants (induced by increasing temperatures, longer growing seasons, and other environmental changes) leads to greater N resorption by trees in autumn, increased C:N in litter, and greater net immobilization of N by soil microbes in the following spring. However, the timing of snowmelt and soil freezing in spring may further affect net mineralization and N availability for plants. These hypotheses are being tested with a combination of observational, experimental, and modeling approaches at Hubbard Brook Experimental Forest in New Hampshire: 1) measurements at 14 previously established sites along an elevation/aspect climate gradient; 2) litter and snow manipulation experiments at six sites along the climate gradient to create variation in soil climate conditions and microbial N immobilization during spring. We leveraged 14 sites previously established along an elevation and aspect-driven climate gradient at Hubbard Brook as a “natural climate experiment" to test our hypothesis that a positive feedback between N cycling during fall senescence and spring contributes to declining N availability in northern forests. This elevation gradient encompasses variation in mean annual air temperature of ~2.5 °C that is similar to the change projected to occur with climate change over the next 50–100 years in the northeastern U.S. There is relatively little variation in soils along the gradient. We are utilizing three sites at higher elevation (~550-660 m, north facing) and three sites at lower elevation (~375-500 m, south facing) for the litter and snow manipulation experiments to maximize the differences in temperature among the 14 sites. Litterbox manipulation: The objective of the litterfall manipulation experiment is to determine whether increases in autumn litter C:N ratios contribute to greater N immobilization by microbes and reductions in net mineralization and plant N uptake in spring, and ultimately, N oligotrophication in northern forest ecosystems. We applied early (low C:N litter that is lost from from hardwood foliage in the first two weeks of autumn) and late (high C:N litter that falls in the last two weeks of autumn) season litter in October 2022 that was collected in fall 2021 at rates equal to standing mass of litter (300 g m2). We also applied native litter that was collected from the forest floor of each intensive site to represent background levels of C:N in litter samples. This litter was applied to one litterbox at each of the six intensive sites. Following application of litter, we installed deer netting around and on top of each of the litterboxes to eliminate litter loss from wind. Soil samples were collected from these plots in November 2021, April 2022, May 2022, June 2022, November 2022, April 2023, May 2023, June 2023 and anlalysed for Nitrogen mineralization and nitrification, as described in the methods section. Snow manipulation: The objective of the snow manipulation experiment is to determine whether the timing of spring snowmelt, length of the spring, and soil freezing in spring affect microbial N immobilization, hydrologic losses, net mineralization, and plant N uptake. The snow manipulation treatment was conducted in the spring of 2022 and 2023. We manually halved (Removal treatment) or doubled (Addition treatment) snow water equivalent (SWE) in experimental plots in March of 2022 and 2023 to accelerate or delay by an average of one week, respectively, the onset of spring snowmelt. Soil Samples were collected from these plots in November 2021, April 2022, May 2022, June 2022, November 2022, April 2023, May 2023, June 2023 and analysed for Nitrogen mineralization and nitrification, as described in the methods section. 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. 

Long-term monitoring of soil nitrate (NO3-) and ammonium (NH4+) concentrations, microbial biomass carbon (C) and nitrogen (N) content, microbial respiration, potential nitrification and N mineralization rates, pH, and denitrification potential has been ongoing at the Hubbard Brook Experimental Forest since 1994. Samples have been collected in the Bear Brook Watershed (west of Watershed 6) beginning in 1994. In 1998, our sampling regime was extended to Watershed 1 in an effort to monitor and quantify microbial response to a whole-watershed calcium addition. 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. 

The goal of this project is to test the overarching hypothesis that positive feedback mechanisms involving changes in seasonal cycles that diminish N availability to plants such that plant N demand is not met by soil N availability in northern forests. Specifically, we hypothesize that increasing N demand by plants (induced by increasing temperatures, longer growing seasons, and other environmental changes) leads to greater N resorption by trees in autumn, increased C:N in litter, and greater net immobilization of N by soil microbes in the following spring. However, the timing of snowmelt and soil freezing in spring may further affect net mineralization and N availability for plants. These hypotheses are being tested with a combination of observational, experimental, and modeling approaches at Hubbard Brook Experimental Forest in New Hampshire: 1) measurements at 14 previously established sites along an elevation/aspect climate gradient; 2) litter and snow manipulation experiments at six sites along the climate gradient to create variation in soil climate conditions and microbial N immobilization during spring. We leveraged 14 sites previously established along an elevation and aspect-driven climate gradient at Hubbard Brook as a “natural climate experiment" to test our hypothesis that a positive feedback between N cycling during fall senescence and spring contributes to declining N availability in northern forests. This elevation gradient encompasses variation in mean annual air temperature of ~2.5 °C that is similar to the change projected to occur with climate change over the next 50–100 years in the northeastern U.S. There is relatively little variation in soils along the gradient. We are utilizing three sites at higher elevation (~550-660 m, north facing) and three sites at lower elevation (~375-500 m, south facing) for the litter and snow manipulation experiments to maximize the differences in temperature among the 14 sites. Litterbox manipulation: The objective of the litterfall manipulation experiment is to determine whether increases in autumn litter C:N ratios contribute to greater N immobilization by microbes and reductions in net mineralization and plant N uptake in spring, and ultimately, N oligotrophication in northern forest ecosystems. We applied early (low C:N litter that is lost from from hardwood foliage in the first two weeks of autumn) and late (high C:N litter that falls in the last two weeks of autumn) season litter in October 2022 that was collected in fall 2021 at rates equal to standing mass of litter (300 g m2). We also applied native litter that was collected from the forest floor of each intensive site to represent background levels of C:N in litter samples. This litter was applied to one litterbox at each of the six intensive sites. Following application of litter, we installed deer netting around and on top of each of the litterboxes to eliminate litter loss from wind. Soil samples were collected from these plots in November 2021, April 2022, May 2022, June 2022, November 2022, April 2023, May 2023, June 2023 and anlalysed for microbial biomass and activity, as described in the methods section. Snow manipulation: The objective of the snow manipulation experiment is to determine whether the timing of spring snowmelt, length of the spring, and soil freezing in spring affect microbial N immobilization, hydrologic losses, net mineralization, and plant N uptake. The snow manipulation treatment was conducted in the spring of 2022 and 2023. We manually halved (Removal treatment) or doubled (Addition treatment) snow water equivalent (SWE) in experimental plots in March of 2022 and 2023 to accelerate or delay by an average of one week, respectively, the onset of spring snowmelt. Soil Samples were collected from these plots in November 2021, April 2022, May 2022, June 2022, November 2022, April 2023, May 2023, June 2023 and anlalysed for microbial biomass and activity, as described in the methods section. 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. 

The Baltimore Ecosystem Study (BES) established a network of long-term permanent biogeochemical study plots in 1998. These plots provide long-term data on vegetation, soil and hydrologic processes in the key ecosystem types within the urban ecosystem. The network of study plots includes forest plots (upland and riparian), chosen to represent the range of forest conditions in the area and grass plots (to represent home lawns). Plots are instrumented with lysimeters (drainage and tension) to sample soil solution chemistry, time domain reflectometry probes to measure soil moisture, dataloggers to measure and record soil temperature, and trace gas flux chambers to measure the flux of carbon dioxide, nitrous oxide and methane from soil to the atmosphere. Measurements of in situ nitrogen mineralization, nitrification and denitrification were made at approximately monthly intervals from Fall 1998 - Fall 2000. Detailed vegetation characterization (all layers) was done in summer 1998 and 2015. Data from these plots has been published in Groffman et al. (2006, 2009), Groffman and Pouyat (2009), Savva et al. (2010), Costa and Groffman (2013), Duncan et al. (2013), Waters et al. (2014), Ni and Groffman (2018), Templeton et al. (2019). Literature Cited Costa, K.H. and P.M. Groffman. 2013. Factors regulating net methane flux in urban forests and grasslands. Soil Science Society of America Journal 77:850 - 855. Duncan, J. M., L. E. Band, and P. M. Groffman. 2013. Towards closing the watershed nitrogen budget: Spatial and temporal scaling of denitrification. Journal of Geophysical Research Biogeosciences 118:1-5; DOI: 10.1002/jgrg.20090 Groffman PM, Pouyat RV, Cadenasso ML, Zipperer WC, Szlavecz K, Yesilonis IC,. Band LE and Brush GS. 2006. Land use context and natural soil controls on plant community composition and soil nitrogen and carbon dynamics in urban and rural forests. Forest Ecology and Management 236:177-192. Groffman, P.M., C.O. Williams, R.V. Pouyat, L.E. Band and I.C. Yesilonis. 2009. Nitrate leaching and nitrous oxide flux in urban forests and grasslands. Journal of Environmental Quality 38:1848-1860. Groffman, P.M. and R.V. Pouyat. 2009. Methane uptake in urban forests and lawns. Environmental Science and Technology 43:5229-5235. DOI: 10.1021/es803720h. Ni, X. and P.M. Groffman. 2018. Declines in methane uptake in forest soils. Proceedings of the National Academies of Science of the United States of America 115:8587-8590. Savva, Y., K. Szlavecz, R. V. Pouyat, P. M. Groffman, and G. Heisler. 2010. Effects of land use and vegetation cover on soil temperature in an urban ecosystem. Soil Science Society of America Journal 74:469-480. Templeton, L., M.L. Cadenasso, J. Sullivan, M. Neel and P.M. Groffman. 2019. Changes in vegetation structure and composition of urban and rural forest patches in Baltimore from 1998 to 2015. Forest Ecology and Management. In press. Waters, E.R., J.L. Morse, N.D. Bettez and P.M. Groffman. 2014. Differential carbon and nitrogen controls of denitrification in riparian zones and streams along an urban to exurban gradient. Journal of Environmental Quality 43:955–963. 

Soil atmosphere fluxes of the trace gases; carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) have been measured at several locations at the Hubbard Brook Experimental Forest (HBEF) including 1) the “freeze” study reference plots that provide contrast between stands dominated (80%) by sugar maple versus yellow birch and low and high elevation areas, 2) the Bear Brook Watershed where trace gas sampling is coordinated with long-term monitoring of microbial biomass and activity and 3) watershed 1 where trace gas sampling locations were co-located with long-term microbial biomass and activity monitoring sites that are located near a subset of the lysimeter sites established for the calcium addition study on this watershed. This dataset contains the Watershed 1 and Bear Brook data. Freeze plot trace gas can be found in: https://portal.edirepository.org/nis/mapbrowse?scope=knb-lter-hbr&identifier=251. 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. 

In the Baltimore urban long-term ecological research (LTER) project, (Baltimore Ecosystem Study, BES) we use the watershed approach to evaluate integrated ecosystem function. The LTER research is centered on the Gwynns Falls watershed, a 17,150 ha catchment that traverses a gradient from the urban core of Baltimore, through older urban residential (1900 - 1950) and suburban (1950- 1980) zones, rapidly suburbanizing areas and a rural/suburban fringe. Our long-term sampling network includes four longitudinal sampling sites along the Gwynns Falls as well as several small (40 - 100 ha) watersheds located within or near to the Gwynns Falls. The longitudinal sites provide data on water and nutrient fluxes in the different land use zones of the watershed (rural/suburban, rapidly suburbanizing, old suburban, urban core) and the small watersheds provide more focused data on specific land use areas (forest, agriculture, rural/suburban, urban). Each of the gaging sites is continuously monitored for discharge and is sampled weekly for chemistry. Additional chemical sampling is carried out in a supplemental set of sites to provide a greater range of land use. Weekly analyses includes nitrate, phosphate, total nitrogen, total phosphorus, chloride and sulfate, turbidity, fecal coliforms, temperature, dissolved oxygen and pH. Cations, dissolved organic carbon and nitrogen and metals are measured on selected samples. Streamflow data for this site are posted at: http://waterdata.usgs.gov/md/nwis/nwisman?site_no=015835701 

The Baltimore Ecosystem Study (BES) established a network of long-term permanent biogeochemical study plots in 1998. These plots provide long-term data on vegetation, soil and hydrologic processes in the key ecosystem types within the urban ecosystem. The network of study plots includes forest plots (upland and riparian), chosen to represent the range of forest conditions in the area and grass plots (to represent home lawns). Plots are instrumented with lysimeters (drainage and tension) to sample soil solution chemistry, time domain reflectometry probes to measure soil moisture, dataloggers to measure and record soil temperature, and trace gas flux chambers to measure the flux of carbon dioxide, nitrous oxide and methane from soil to the atmosphere. Measurements of in situ nitrogen mineralization, nitrification and denitrification were made at approximately monthly intervals from Fall 1998 - Fall 2000. Detailed vegetation characterization (all layers) was done in summer 1998 and 2015. Data from these plots has been published in Groffman et al. (2006, 2009), Groffman and Pouyat (2009), Savva et al. (2010), Costa and Groffman (2013), Duncan et al. (2013), Waters et al. (2014), Ni and Groffman (2018), Templeton et al. (2019). Literature Cited Costa, K.H. and P.M. Groffman. 2013. Factors regulating net methane flux in urban forests and grasslands. Soil Science Society of America Journal 77:850 - 855. Duncan, J. M., L. E. Band, and P. M. Groffman. 2013. Towards closing the watershed nitrogen budget: Spatial and temporal scaling of denitrification. Journal of Geophysical Research Biogeosciences 118:1-5; DOI: 10.1002/jgrg.20090 Groffman PM, Pouyat RV, Cadenasso ML, Zipperer WC, Szlavecz K, Yesilonis IC,. Band LE and Brush GS. 2006. Land use context and natural soil controls on plant community composition and soil nitrogen and carbon dynamics in urban and rural forests. Forest Ecology and Management 236:177-192. Groffman, P.M., C.O. Williams, R.V. Pouyat, L.E. Band and I.C. Yesilonis. 2009. Nitrate leaching and nitrous oxide flux in urban forests and grasslands. Journal of Environmental Quality 38:1848-1860. Groffman, P.M. and R.V. Pouyat. 2009. Methane uptake in urban forests and lawns. Environmental Science and Technology 43:5229-5235. DOI: 10.1021/es803720h. Ni, X. and P.M. Groffman. 2018. Declines in methane uptake in forest soils. Proceedings of the National Academies of Science of the United States of America 115:8587-8590. Savva, Y., K. Szlavecz, R. V. Pouyat, P. M. Groffman, and G. Heisler. 2010. Effects of land use and vegetation cover on soil temperature in an urban ecosystem. Soil Science Society of America Journal 74:469-480. Templeton, L., M.L. Cadenasso, J. Sullivan, M. Neel and P.M. Groffman. 2019. Changes in vegetation structure and composition of urban and rural forest patches in Baltimore from 1998 to 2015. Forest Ecology and Management. In press. Waters, E.R., J.L. Morse, N.D. Bettez and P.M. Groffman. 2014. Differential carbon and nitrogen controls of denitrification in riparian zones and streams along an urban to exurban gradient. Journal of Environmental Quality 43:955–963. 

Soil atmosphere fluxes of the trace gases; carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) have been measured at several locations at the Hubbard Brook Experimental Forest (HBEF) including 1) the “freeze” study reference plots that provide contrast between stands dominated (80%) by sugar maple versus yellow birch and low and high elevation areas, 2) the Bear Brook Watershed where trace gas sampling is coordinated with long-term monitoring of microbial biomass and activity and 3) watershed 1 where trace gas sampling locations were co-located with long-term microbial biomass and activity monitoring sites that are located near a subset of the lysimeter sites established for the calcium addition study on this watershed. This dataset contains the Freeze study data. Watershed 1 and Bear Brook trace gas data can be found in: https://portal.edirepository.org/nis/mapbrowse?scope=knb-lter-hbr&identifier=116. 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 published in: Groffman, P. M., Hardy, J. P., Driscoll, C. T., & Fahey, T. J. (2006). Snow depth, soil freezing, and fluxes of carbon dioxide, nitrous oxide and methane in a northern hardwood forest. Global Change Biology, 12, 1748–1760. 

Soil atmosphere fluxes of the trace gases; carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) have been measured at several locations at the Hubbard Brook Experimental Forest (HBEF) including 1) the “freeze” study reference plots that provide contrast between stands dominated (80%) by sugar maple versus yellow birch and low and high elevation areas, 2) the Bear Brook Watershed where trace gas sampling is coordinated with long-term monitoring of microbial biomass and activity and 3) watershed 1 where trace gas sampling locations were co-located with long-term microbial biomass and activity monitoring sites that are located near a subset of the lysimeter sites established for the calcium addition study on this watershed. This dataset contains the Watershed 1 and Bear Brook data. Freeze plot trace gas can be found in: https://portal.edirepository.org/nis/mapbrowse?scope=knb-lter-hbr&identifier=251. 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. 

Long-term monitoring of soil nitrate (NO3-) and ammonium (NH4+) concentrations, microbial biomass carbon (C) and nitrogen (N) content, microbial respiration, potential nitrification and N mineralization rates, pH, and denitrification potential has been ongoing at the Hubbard Brook Experimental Forest since 1994. Samples have been collected in the Bear Brook Watershed (west of Watershed 6) beginning in 1994. In 1998, our sampling regime was extended to Watershed 1 in an effort to monitor and quantify microbial response to a whole-watershed calcium addition. 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.