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• 1. [Article] Impacts of forest-to-agriculture conversion on aboveground and soil carbon and nitrogen stocks along a bioclimatic gradient in Costa Rica

  Tropical forest-to-agriculture conversions contribute to ~20% of contemporary global C emissions. Current global C flux estimates assume C is lost from vegetation and soils over time due to this conversion, ...

  Citation × Citation Citation Abstract Copy Title: Impacts of forest-to-agriculture conversion on aboveground and soil carbon and nitrogen stocks along a bioclimatic gradient in Costa Rica Author: Jobse, Judith C. Tropical forest-to-agriculture conversions contribute to ~20% of contemporary global C emissions. Current global C flux estimates assume C is lost from vegetation and soils over time due to this conversion, and that C stocks in agricultural lands are fairly uniform across the tropics. Global C stock and flux estimates may contain large errors if these assumptions are incorrect. I asked the following research questions: (1) Do aboveground and soil C and N stocks and their net changes due to forest-to-agricultural conversion differ among life zones and agricultural land use types in Costa Rica?; (2) Do aboveground and soil C and N stocks and their net changes due to forest-to-pasture conversion differ by pasture age?; and (3) Can life zone specific-estimates reduce potential biases and uncertainty of regional and global C flux estimates for forest-to-agricultural conversions? I sampled soil and aboveground C and N stocks at 62 agricultural sites in Costa Rica stratified by land use and life zone and compared them to mature forests. Ecosystem and soil C and N stocks varied more by life zone than by land use. Net soil C and N stock changes due to forest-to-pasture conversion differed by life zone; soil stocks decreased in the Tropical dry forest and Lower Montane rain forest life zones and increased in the other life zones. Generally, ecosystem C and N stocks in pastures decreased with pasture age, but the effect of age was smaller than life zone and conversion effects. My estimates of Costa Rican ecosystem C stocks in agricultural lands were 2-fold greater and my ecosystem C flux estimates due to forest-to-agricultural conversions were 8-19% lower than Intergovernmental Panel on Climate Change (IPCC)-endorsed estimates. Therefore to reduce uncertainty and bias, incorporation of ecosystem C stock variability related to life zone into regional and global models is more important than including variability related to pasture age or land use type. Overall, my results confirm that forest-to-agriculture conversions represent a significant alteration to global C cycles, and contribute to reducing uncertainty in the magnitude of such changes. Title: Impacts of forest-to-agriculture conversion on aboveground and soil carbon and nitrogen stocks along a bioclimatic gradient in Costa Rica Author: Jobse, Judith C. Tropical forest-to-agriculture conversions contribute to ~20% of contemporary global C emissions. Current global C flux estimates assume C is lost from vegetation and soils over time due to this conversion, and that C stocks in agricultural lands are fairly uniform across the tropics. Global C stock and flux estimates may contain large errors if these assumptions are incorrect. I asked the following research questions: (1) Do aboveground and soil C and N stocks and their net changes due to forest-to-agricultural conversion differ among life zones and agricultural land use types in Costa Rica?; (2) Do aboveground and soil C and N stocks and their net changes due to forest-to-pasture conversion differ by pasture age?; and (3) Can life zone specific-estimates reduce potential biases and uncertainty of regional and global C flux estimates for forest-to-agricultural conversions? I sampled soil and aboveground C and N stocks at 62 agricultural sites in Costa Rica stratified by land use and life zone and compared them to mature forests. Ecosystem and soil C and N stocks varied more by life zone than by land use. Net soil C and N stock changes due to forest-to-pasture conversion differed by life zone; soil stocks decreased in the Tropical dry forest and Lower Montane rain forest life zones and increased in the other life zones. Generally, ecosystem C and N stocks in pastures decreased with pasture age, but the effect of age was smaller than life zone and conversion effects. My estimates of Costa Rican ecosystem C stocks in agricultural lands were 2-fold greater and my ecosystem C flux estimates due to forest-to-agricultural conversions were 8-19% lower than Intergovernmental Panel on Climate Change (IPCC)-endorsed estimates. Therefore to reduce uncertainty and bias, incorporation of ecosystem C stock variability related to life zone into regional and global models is more important than including variability related to pasture age or land use type. Overall, my results confirm that forest-to-agriculture conversions represent a significant alteration to global C cycles, and contribute to reducing uncertainty in the magnitude of such changes. Close

• 2. [Article] Four centuries of soil carbon and nitrogen change after severe fire in a Western Cascades forest landscape

  Fire is a major disturbance process in many forests. Long-term studies of the biogeochemical effects of fires, especially on soils, are very rare. Consequently, long-term effects of fire on soils are often ...

  Citation × Citation Citation Abstract Copy Title: Four centuries of soil carbon and nitrogen change after severe fire in a Western Cascades forest landscape Author: Giesen, Thomas William Fire is a major disturbance process in many forests. Long-term studies of the biogeochemical effects of fires, especially on soils, are very rare. Consequently, long-term effects of fire on soils are often hypothesized from short-term effects. In a chronosequence study, I studied 24 western Cascades (Oregon) forest stands thought to have been initiated in fire. Twelve of those burned about 150 years ago (“young” sites), and the other 12 burned an average of 550 years ago (“old” sites). I hypothesized that young stands would have less carbon (C) and nitrogen (N) in forest floor and in 0 -10 cm mineral soil than old stands. I found that forest floor N pools of old sites (average = 1,823 kg/ha ± s.e. = 132 kg/ha) were significantly greater than young sites (1,450 ± 98 kg/ha). Similarly, forest floor C pools of old sites (62,980 ± 5,403 kg/ha) were significantly greater than young sites (49,032 ± 2,965 kg/ha). Greater N and C pools in forest floor of old sites resulted from greater forest floor mass in old sites; concentrations of both N and C, and C:N ratios, did not differ significantly by forest age class. In mineral soil, neither concentrations nor pools of N and C differed between young and old sites. Despite overall similarity of C:N ratios in young versus old sites, potential N mineralization rates were twice as high in forest floor of old sites (average = 60 ± 7.3 mg N / g soil) than young sites (26 ± 3.5 mg N / g soil), . Nitrate accounted for only 2% or less of total N mineralized in forest floor samples. In mineral soil, potential net N mineralization did not differ by forest age class. The pattern of high net N mineralization and low nitrification in old forests is consistent with other studies of fire-prone forests, yet contrasts with many studies of forests that lack fire, and suggests that ammonium is not the sole control over nitrification in fire-prone ecosystems. Overall, fire appears to impart a longterm legacy of reduced forest floor N and C pools in this western Oregon Cascades landscape, which suggests that current fire-suppression activities in the region may increase forest floor N and C storage over historical conditions within several centuries. The differences in forest floor and soil N cycling processes that I observed by forest age class raise the further possibility that fire exclusion in these forests may change the relative abundance of soil inorganic N forms to favor ammonium over nitrate. Such changes may have unknown consequences for relative competitive abilities of plant and microbial species that rely preferentially on different N-forms to meet N nutrition requirements. While forest floor N and C pools increase from young to old stands, forest floor and soil N and C pools are not different, or decline, between 450 year old stands and the oldest stands at 800+ years, That, and other, anomalous changes in values from ~450 to 800+ years, suggest possible changes in ecosystem functions, and may indicate that this landscape could be a fruitful study area for examinations of a mature, steady-state ecosystem Title: Four centuries of soil carbon and nitrogen change after severe fire in a Western Cascades forest landscape Author: Giesen, Thomas William Fire is a major disturbance process in many forests. Long-term studies of the biogeochemical effects of fires, especially on soils, are very rare. Consequently, long-term effects of fire on soils are often hypothesized from short-term effects. In a chronosequence study, I studied 24 western Cascades (Oregon) forest stands thought to have been initiated in fire. Twelve of those burned about 150 years ago (“young” sites), and the other 12 burned an average of 550 years ago (“old” sites). I hypothesized that young stands would have less carbon (C) and nitrogen (N) in forest floor and in 0 -10 cm mineral soil than old stands. I found that forest floor N pools of old sites (average = 1,823 kg/ha ± s.e. = 132 kg/ha) were significantly greater than young sites (1,450 ± 98 kg/ha). Similarly, forest floor C pools of old sites (62,980 ± 5,403 kg/ha) were significantly greater than young sites (49,032 ± 2,965 kg/ha). Greater N and C pools in forest floor of old sites resulted from greater forest floor mass in old sites; concentrations of both N and C, and C:N ratios, did not differ significantly by forest age class. In mineral soil, neither concentrations nor pools of N and C differed between young and old sites. Despite overall similarity of C:N ratios in young versus old sites, potential N mineralization rates were twice as high in forest floor of old sites (average = 60 ± 7.3 mg N / g soil) than young sites (26 ± 3.5 mg N / g soil), . Nitrate accounted for only 2% or less of total N mineralized in forest floor samples. In mineral soil, potential net N mineralization did not differ by forest age class. The pattern of high net N mineralization and low nitrification in old forests is consistent with other studies of fire-prone forests, yet contrasts with many studies of forests that lack fire, and suggests that ammonium is not the sole control over nitrification in fire-prone ecosystems. Overall, fire appears to impart a longterm legacy of reduced forest floor N and C pools in this western Oregon Cascades landscape, which suggests that current fire-suppression activities in the region may increase forest floor N and C storage over historical conditions within several centuries. The differences in forest floor and soil N cycling processes that I observed by forest age class raise the further possibility that fire exclusion in these forests may change the relative abundance of soil inorganic N forms to favor ammonium over nitrate. Such changes may have unknown consequences for relative competitive abilities of plant and microbial species that rely preferentially on different N-forms to meet N nutrition requirements. While forest floor N and C pools increase from young to old stands, forest floor and soil N and C pools are not different, or decline, between 450 year old stands and the oldest stands at 800+ years, That, and other, anomalous changes in values from ~450 to 800+ years, suggest possible changes in ecosystem functions, and may indicate that this landscape could be a fruitful study area for examinations of a mature, steady-state ecosystem Close

• 3. [Article] Non-linear nitrogen dynamics and calcium depletion along a temperate forest soil nitrogen gradient

  Understanding how N availability influences base cation stores is critical for long-term ecosystem sustainability. Indices of nitrogen (N) availability and the distribution of nutrients in plant biomass, ...

  Citation × Citation Citation Abstract Copy Title: Non-linear nitrogen dynamics and calcium depletion along a temperate forest soil nitrogen gradient Author: Sinkhorn, Emily R. Understanding how N availability influences base cation stores is critical for long-term ecosystem sustainability. Indices of nitrogen (N) availability and the distribution of nutrients in plant biomass, soil, and soil water were examined across ten young, unpolluted Douglas-fir (Pseudotsuga menziesii) stands in the Oregon Coast Range spanning a three-fold soil N gradient (0-10 cm: 0.21 - 0.69% N, 0-100 cm: 9.2 – 28.8 Mg N . ha⁻¹) but having similar stand age and sandstone parent material. δ¹⁵N in foliage and forest floor increased across the gradient and approached the isotopic signature of the atmosphere at high soil N stands, suggesting that variation in N accumulation across sites is related to historic site occupancy by N₂-fixing red alder (Alnus rubra). Although no longer present on these sites, red alder stands can add 100-200 kg N ha⁻¹ yr⁻¹ to an ecosystem for decades, a significantly higher N input than precipitation (0.65 kg N ha⁻¹ yr⁻¹). Annual net N mineralization and litterfall N return displayed non-linear relationships with soil N, increasing initially, and then decreasing at more N-rich sites. In contrast, nitrate leaching from deep soils increased linearly across the soil N gradient and ranged from 0.074 to 30 kg N . ha⁻¹ . yr⁻¹. Nitrogen availability was negatively correlated with indices of Ca availability. Soil exchangeable Ca, Mg, and K pools to 1 m depth were negatively related to nitrate losses across sites. Calcium was the only base cation that decreased in both plant and soil pools across the soil N gradient, and a greater proportion of total available ecosystem Ca was sequestered in plant biomass at high N, low Ca sites. The preferential storage of Ca in aboveground biomass at high N and low Ca sites, while critical for sustaining plant productivity, may also predispose forests to Ca depletion in areas managed for intensive biomass removal. Our work supports a hierarchical model of coupled N-Ca cycles across gradients of soil N enrichment, with microbial production of mobile nitrate leading to depletion of readily available Ca at the ecosystem scale, and plant sequestration promoting Ca conservation as Ca supply diminishes. Long-term N enrichment of temperate forest soils appears capable of sustaining an open N cycle and key symptoms of N saturation for multiple decades after the cessation of elevated N inputs. Title: Non-linear nitrogen dynamics and calcium depletion along a temperate forest soil nitrogen gradient Author: Sinkhorn, Emily R. Understanding how N availability influences base cation stores is critical for long-term ecosystem sustainability. Indices of nitrogen (N) availability and the distribution of nutrients in plant biomass, soil, and soil water were examined across ten young, unpolluted Douglas-fir (Pseudotsuga menziesii) stands in the Oregon Coast Range spanning a three-fold soil N gradient (0-10 cm: 0.21 - 0.69% N, 0-100 cm: 9.2 – 28.8 Mg N . ha⁻¹) but having similar stand age and sandstone parent material. δ¹⁵N in foliage and forest floor increased across the gradient and approached the isotopic signature of the atmosphere at high soil N stands, suggesting that variation in N accumulation across sites is related to historic site occupancy by N₂-fixing red alder (Alnus rubra). Although no longer present on these sites, red alder stands can add 100-200 kg N ha⁻¹ yr⁻¹ to an ecosystem for decades, a significantly higher N input than precipitation (0.65 kg N ha⁻¹ yr⁻¹). Annual net N mineralization and litterfall N return displayed non-linear relationships with soil N, increasing initially, and then decreasing at more N-rich sites. In contrast, nitrate leaching from deep soils increased linearly across the soil N gradient and ranged from 0.074 to 30 kg N . ha⁻¹ . yr⁻¹. Nitrogen availability was negatively correlated with indices of Ca availability. Soil exchangeable Ca, Mg, and K pools to 1 m depth were negatively related to nitrate losses across sites. Calcium was the only base cation that decreased in both plant and soil pools across the soil N gradient, and a greater proportion of total available ecosystem Ca was sequestered in plant biomass at high N, low Ca sites. The preferential storage of Ca in aboveground biomass at high N and low Ca sites, while critical for sustaining plant productivity, may also predispose forests to Ca depletion in areas managed for intensive biomass removal. Our work supports a hierarchical model of coupled N-Ca cycles across gradients of soil N enrichment, with microbial production of mobile nitrate leading to depletion of readily available Ca at the ecosystem scale, and plant sequestration promoting Ca conservation as Ca supply diminishes. Long-term N enrichment of temperate forest soils appears capable of sustaining an open N cycle and key symptoms of N saturation for multiple decades after the cessation of elevated N inputs. Close

• 4. [Article] Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon

  Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. I investigated patterns ...

  Citation × Citation Citation Abstract Copy Title: Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon Author: Thiel, Aaron L. Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. I investigated patterns of nitrogen (N) availability along transects through 0.1 and 0.4 ha silvicultural gaps in three 50-70 year old Douglas-fir forests of western Oregon. Six indices of N availability in forest floor and mineral soil and several factors related to N cycling were measured from November 2005 to February 2007, approximately 6-8 years after gap creation. Results indicate that mineral soil pools of extractable ammonium (NH4⁺) and nitrate (NO3⁻), rates of net N mineralization and nitrification, and concentrations of ion-exchange resin NH4⁺ and NO3⁻ were significantly elevated in gaps relative to adjacent forest. Gap-forest differences in forest floor layers were less clear. For the majority of response variables, magnitudes and trends were similar in the centers of both gap sizes. N availability in gap edge positions more often resembled levels in the forest than in the gap interior, and there were few significant differences between positions north and south of gap centers. Forest floor and mineral soil percent moisture did not significantly differ along gap transects, nor did decomposition rates of wooden tongue depressors. Litterfall carbon (C) inputs and litterfall C:N ratios in gaps were significantly lower than in the forest. Reciprocal transfer incubations of mineral soil samples between gap and forest positions revealed that sample origin had a significant effect on net nitrification rates, while incubation environment did not. Variability of several indices of N availability also increased in gaps. The overall increase of N availability in 6-8 year old silvicultural gaps may be due more to the quality and quantity of litterfall inputs than temperature and moisture conditions. Increased quality of litterfall in gaps, as indicated by lower C:N ratios, may increase rates of decomposition and net N mineralization, while overall lower litterfall C inputs may lead to C-limitation of microbial immobilization, resulting in increased accumulation of inorganic N in soil. While environmental factors have been shown to drive N availability soon after gap creation, litter inputs from early-seral species may perpetuate increased N availability into early stages of vegetative succession. From a management perspective, increased N availability in gaps may increase tree productivity, but at the same time, increase the likelihood of invasion by exotic species. Gap creation may also increase gap-scale heterogeneity of available N in the short-term, while increasing stand-scale heterogeneity in the long-term. Title: Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon Author: Thiel, Aaron L. Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. I investigated patterns of nitrogen (N) availability along transects through 0.1 and 0.4 ha silvicultural gaps in three 50-70 year old Douglas-fir forests of western Oregon. Six indices of N availability in forest floor and mineral soil and several factors related to N cycling were measured from November 2005 to February 2007, approximately 6-8 years after gap creation. Results indicate that mineral soil pools of extractable ammonium (NH4⁺) and nitrate (NO3⁻), rates of net N mineralization and nitrification, and concentrations of ion-exchange resin NH4⁺ and NO3⁻ were significantly elevated in gaps relative to adjacent forest. Gap-forest differences in forest floor layers were less clear. For the majority of response variables, magnitudes and trends were similar in the centers of both gap sizes. N availability in gap edge positions more often resembled levels in the forest than in the gap interior, and there were few significant differences between positions north and south of gap centers. Forest floor and mineral soil percent moisture did not significantly differ along gap transects, nor did decomposition rates of wooden tongue depressors. Litterfall carbon (C) inputs and litterfall C:N ratios in gaps were significantly lower than in the forest. Reciprocal transfer incubations of mineral soil samples between gap and forest positions revealed that sample origin had a significant effect on net nitrification rates, while incubation environment did not. Variability of several indices of N availability also increased in gaps. The overall increase of N availability in 6-8 year old silvicultural gaps may be due more to the quality and quantity of litterfall inputs than temperature and moisture conditions. Increased quality of litterfall in gaps, as indicated by lower C:N ratios, may increase rates of decomposition and net N mineralization, while overall lower litterfall C inputs may lead to C-limitation of microbial immobilization, resulting in increased accumulation of inorganic N in soil. While environmental factors have been shown to drive N availability soon after gap creation, litter inputs from early-seral species may perpetuate increased N availability into early stages of vegetative succession. From a management perspective, increased N availability in gaps may increase tree productivity, but at the same time, increase the likelihood of invasion by exotic species. Gap creation may also increase gap-scale heterogeneity of available N in the short-term, while increasing stand-scale heterogeneity in the long-term. Close

• 5. [Article] Interactions between ecosystem nitrogen and bedrock control long-term calcium sources in Oregon Coast Range forests

  Ecosystem nitrogen (N) supply strongly influences the availability and cycling of other essential nutrients in temperate forests, especially calcium (Ca). Short-term additions of N that exceed ecosystem ...

  Citation × Citation Citation Abstract Copy Title: Interactions between ecosystem nitrogen and bedrock control long-term calcium sources in Oregon Coast Range forests Author: Hynicka, Justin D. Ecosystem nitrogen (N) supply strongly influences the availability and cycling of other essential nutrients in temperate forests, especially calcium (Ca). Short-term additions of N that exceed ecosystem demands often increase dissolved nitrate fluxes and decrease soil pH, which can stimulate soil Ca loss. However, the long-term effects of high N supply on ecosystem Ca availability are more difficult to determine, and may depend on the Ca content of bedrock and mineral soils. To address this, we examined major and trace element concentrations and ⁸⁷Sr/⁸⁶Sr ratios that trace Ca sources in precipitation, foliage, soil pools, and bedrock at 24 forested sites in the Oregon Coast Range having a wide, natural range of soil N (0.16 - 0.97 % N, 0-10 cm) on contrasting basaltic and sedimentary bedrock. Using a suite of 17 site properties, we also evaluated whether soil N variation across sites was related to the five major state-factors of soil and ecosystem development: climate, organisms, topography, parent material, and time. We found that as soil N increased across sites, its ¹⁵N/¹⁴N ratio declined towards atmospheric values, suggesting that soil N variation reflects a biotic legacy of symbiotic N fixation inputs. In contrast, soil N variation was unrelated to 17 other metrics of soil forming factors that represented climate (mean annual precipitation, mean annual temperature, and distance from the coast), topography (slope, soil depth, and abundance of coarse rock fragments), parent material (within bedrock type bulk and 1 M HNO₃ leachable rock Ca chemistry), and proxies of soil age (Hurst's redness rating, effective cation exchange capacity, Ca in non-exchangeable soil residues, chemical index of alteration, weathering index of Parker, Ca in coarse soil fragments, and soil Ca loss relative to bedrock). These analyses highlight symbiotic N-fixing red alder as a keystone organismal state-factor that produces a wide range of soil N accumulation in coastal Oregon forests. Strontium isotopes (⁸⁷Sr/⁸⁶Sr) and other geochemical analyses indicate that long-term Ca sources in foliage and exchangeable soil pools in Oregon Coast Range forests depend on an interactive effect between N availability and bedrock. Basaltic rocks contained nearly 20-times more Ca than sedimentary rocks across our sites, and this difference was reflected in Sr-isotope partitioning of base cation sources. Atmospheric sources dominated plant and soil pools in forests overlying Ca-poor sedimentary rock, regardless of variation in soil N, indicating extremely limited capacity of weathering to support forest Ca demands. In contrast, forests overlying basaltic rock obtained as much as 80% of Ca from rock weathering in low N sites, yet relied to a greater extent on atmospheric Ca as soil N increased, with less than 10% of Ca from rock weathering at sites with the highest soil N. Surprisingly, differences in fresh rock Ca content and base cation sources between sedimentary and basaltic sites was not reflected in ecosystem Ca availability, and instead increasing soil N caused similar declines in foliar and exchangeable Ca across both rock types. This illustrates that nutrient pool sizes do not necessarily reflect long-term nutrient supply, and highlights how coupled biogeochemical cycles within ecosystems can regulate nutrient loss and supply to biota. Broadly, our results highlight how interactions between biological and geologic factors can influence base cation sources in forest ecosystems. The sustainability of base cation supplies to forests may therefore depend greatly on variation in bedrock weathering at low N sites, yet converge to depend on atmospheric inputs in sites that receive high N loading from biological fixation or anthropogenic deposition. Title: Interactions between ecosystem nitrogen and bedrock control long-term calcium sources in Oregon Coast Range forests Author: Hynicka, Justin D. Ecosystem nitrogen (N) supply strongly influences the availability and cycling of other essential nutrients in temperate forests, especially calcium (Ca). Short-term additions of N that exceed ecosystem demands often increase dissolved nitrate fluxes and decrease soil pH, which can stimulate soil Ca loss. However, the long-term effects of high N supply on ecosystem Ca availability are more difficult to determine, and may depend on the Ca content of bedrock and mineral soils. To address this, we examined major and trace element concentrations and ⁸⁷Sr/⁸⁶Sr ratios that trace Ca sources in precipitation, foliage, soil pools, and bedrock at 24 forested sites in the Oregon Coast Range having a wide, natural range of soil N (0.16 - 0.97 % N, 0-10 cm) on contrasting basaltic and sedimentary bedrock. Using a suite of 17 site properties, we also evaluated whether soil N variation across sites was related to the five major state-factors of soil and ecosystem development: climate, organisms, topography, parent material, and time. We found that as soil N increased across sites, its ¹⁵N/¹⁴N ratio declined towards atmospheric values, suggesting that soil N variation reflects a biotic legacy of symbiotic N fixation inputs. In contrast, soil N variation was unrelated to 17 other metrics of soil forming factors that represented climate (mean annual precipitation, mean annual temperature, and distance from the coast), topography (slope, soil depth, and abundance of coarse rock fragments), parent material (within bedrock type bulk and 1 M HNO₃ leachable rock Ca chemistry), and proxies of soil age (Hurst's redness rating, effective cation exchange capacity, Ca in non-exchangeable soil residues, chemical index of alteration, weathering index of Parker, Ca in coarse soil fragments, and soil Ca loss relative to bedrock). These analyses highlight symbiotic N-fixing red alder as a keystone organismal state-factor that produces a wide range of soil N accumulation in coastal Oregon forests. Strontium isotopes (⁸⁷Sr/⁸⁶Sr) and other geochemical analyses indicate that long-term Ca sources in foliage and exchangeable soil pools in Oregon Coast Range forests depend on an interactive effect between N availability and bedrock. Basaltic rocks contained nearly 20-times more Ca than sedimentary rocks across our sites, and this difference was reflected in Sr-isotope partitioning of base cation sources. Atmospheric sources dominated plant and soil pools in forests overlying Ca-poor sedimentary rock, regardless of variation in soil N, indicating extremely limited capacity of weathering to support forest Ca demands. In contrast, forests overlying basaltic rock obtained as much as 80% of Ca from rock weathering in low N sites, yet relied to a greater extent on atmospheric Ca as soil N increased, with less than 10% of Ca from rock weathering at sites with the highest soil N. Surprisingly, differences in fresh rock Ca content and base cation sources between sedimentary and basaltic sites was not reflected in ecosystem Ca availability, and instead increasing soil N caused similar declines in foliar and exchangeable Ca across both rock types. This illustrates that nutrient pool sizes do not necessarily reflect long-term nutrient supply, and highlights how coupled biogeochemical cycles within ecosystems can regulate nutrient loss and supply to biota. Broadly, our results highlight how interactions between biological and geologic factors can influence base cation sources in forest ecosystems. The sustainability of base cation supplies to forests may therefore depend greatly on variation in bedrock weathering at low N sites, yet converge to depend on atmospheric inputs in sites that receive high N loading from biological fixation or anthropogenic deposition. Close