Patterns of Nutrient Loss from Unpolluted, Old-Growth Temperate Forests: Evaluation of Biogeochemical Theory

Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find r...

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Published inEcology (Durham) Vol. 76; no. 2; pp. 493 - 509
Main Authors Hedin, Lars O., Armesto, Juan J., Johnson, Arthur H.
Format Journal Article
LanguageEnglish
Published Washington, DC The Ecological Society of America 01.03.1995
Ecological Society of America
Brooklyn Botanic Garden, etc
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Abstract Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important @?baseline@? information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional-scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the @?nutrient retention hypothesis@?) that net biotic retention of elements should be minimal in old-growth ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrolic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO"3 ^- (0.2% of total N) and NH"4^+ (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old-growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long-term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old-growth forests as @?leaky@? vs. @?non-leaky@? with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High- and mid-elevation forests in our study area showed the lowest afflux concentrations of N as NO"3 - reported from any Old-growth temperate forest ecosystem (0.l0 vs. 0.30 @mg/L, respectively). Comparisons against other old-growth forests, subject to varying levels of N deposition, indicated that levels of NO"3^- and the relative abundance of NO"3^- vs. NH"4^+ in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old-growth forest ecosystems are particularly sensitive indicators of N deposition.
AbstractList The patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds are presented.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important °baseline° information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional—scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old—growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the °nutrient retention hypothesis°) that net biotic retention of elements should be minimal in old—growth ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrolic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO 3 — (0.2% of total N) and NH 4 + (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old—growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long—term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old—growth forests as °leaky° vs. °non—leaky° with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High— and mid—elevation forests in our study area showed the lowest afflux concentrations of N as NO 3 — reported from any Old—growth temperate forest ecosystem (0.l0 vs. 0.30 mg/L, respectively). Comparisons against other old—growth forests, subject to varying levels of N deposition, indicated that levels of NO 3 — and the relative abundance of NO 3 — vs. NH 4 + in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old—growth forest ecosystems are particularly sensitive indicators of N deposition.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important "baseline" information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional-scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the "nutrient retention hypothesis") that net biotic retention of elements should be minimal in old-growth forest ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrologic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO sub(3) super(-) (0.2% of total N) and NH sub(4) super(+) (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old-growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long-term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old-growth forests as "leaky" vs. "non-leaky" with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High- and mid-elevation forests in our study area showed the lowest efflux concentrations of N as NO sub(3) super(-) reported from any old-growth temperate forest ecosystem (0.10 vs. 0.30 mu g/L, respectively). Comparisons against other old-growth forests, subject to varying levels of N deposition, indicated that levels of NO sub(3) super(-) and the relative abundance of NO sub(3) super(-) vs. NH sub(4) super(+) in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional-scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old-growth forest ecosystems are particularly sensitive indicators of N deposition.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important °baseline° information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional—scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old—growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the °nutrient retention hypothesis°) that net biotic retention of elements should be minimal in old—growth ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrolic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO3 — (0.2% of total N) and NH4+ (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old—growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long—term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old—growth forests as °leaky° vs. °non—leaky° with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High— and mid—elevation forests in our study area showed the lowest afflux concentrations of N as NO3 — reported from any Old—growth temperate forest ecosystem (0.l0 vs. 0.30 mg/L, respectively). Comparisons against other old—growth forests, subject to varying levels of N deposition, indicated that levels of NO3— and the relative abundance of NO3— vs. NH4+ in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old—growth forest ecosystems are particularly sensitive indicators of N deposition.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas, While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important "baseline" information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional-scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the "nutrient retention hypothesis") that net biotic retention of elements should be minimal in old-growth forest ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrologic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms N[O.sub.[3.sup.-]] (0.2% of total N) and N[H.sub.[4.sup.+]] (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old-growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long-term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old-growth forests as "leaky" vs. "non-leaky" with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High- and mid-elevation forests in our study area showed the lowest efflux concentrations of N as N[O.sub.[3.sup.-]] reported from any old-growth temperate forest ecosystem (0.10 vs. 0.30 [µgram]/L, respectively). Comparisons against other old-growth forests, subject to varying levels of N deposition, indicated that levels of N[O.sub.[3.sup.-]] and the relative abundance of N[O.sub.[3.sup.-]] vs. N[H.sub.[4.sup.+]] in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional-scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old-growth forest ecosystems are particularly sensitive indicators of N deposition. Key words: atmospheric deposition; base cations; biogeochemical theory; biogeochemistry; Chile; nitrogen; nitrogen saturation hypothesis; nutrient cycling; nutrient retention hypothesis; old-growth forests; temperate forest ecosystems; unpolluted conditions; watershed approach.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important °baseline° information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional—scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old—growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the °nutrient retention hypothesis°) that net biotic retention of elements should be minimal in old—growth ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrolic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO₃ — (0.2% of total N) and NH₄ ⁺ (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old—growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long—term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old—growth forests as °leaky° vs. °non—leaky° with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High— and mid—elevation forests in our study area showed the lowest afflux concentrations of N as NO₃ — reported from any Old—growth temperate forest ecosystem (0.l0 vs. 0.30 mg/L, respectively). Comparisons against other old—growth forests, subject to varying levels of N deposition, indicated that levels of NO₃ — and the relative abundance of NO₃ — vs. NH₄ ⁺ in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old—growth forest ecosystems are particularly sensitive indicators of N deposition.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas. While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important @?baseline@? information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional-scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the @?nutrient retention hypothesis@?) that net biotic retention of elements should be minimal in old-growth ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrolic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms NO"3 ^- (0.2% of total N) and NH"4^+ (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old-growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long-term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old-growth forests as @?leaky@? vs. @?non-leaky@? with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High- and mid-elevation forests in our study area showed the lowest afflux concentrations of N as NO"3 - reported from any Old-growth temperate forest ecosystem (0.l0 vs. 0.30 @mg/L, respectively). Comparisons against other old-growth forests, subject to varying levels of N deposition, indicated that levels of NO"3^- and the relative abundance of NO"3^- vs. NH"4^+ in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old-growth forest ecosystems are particularly sensitive indicators of N deposition.
Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient dynamics in temperate region ecosystems has been derived from such biogeochemically altered areas, While it is increasingly difficult to find regions of temperate forest that can be classified as free from human influences, studies of unpolluted and undisturbed areas can (1) provide important "baseline" information about natural patterns of element cycling, against which disturbed cycles can be compared; (2) provide insights into the biogeochemical conditions that acted as evolutionary and selective constraints on biotic communities before the advent of regional-scale human impacts; and (3) provide an opportunity to evaluate the general nature of biogeochemical theories that have been developed in areas subject to strong human influences. We here report on patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31 watersheds. These forests have not been subject to air pollution inputs and have remained floristically stable throughout the Holocene. For major elements our results support predictions from current biogeochemical theory (the "nutrient retention hypothesis") that net biotic retention of elements should be minimal in old-growth forest ecosystems. Despite the dilute nature of watershed streams, the overall patterns of loss of most elements could be explained almost exclusively by atmospheric inputs of marine aerosols. In contrast, patterns of nitrogen loss were more complex than predicted by current conceptual models. Hydrologic N losses occurred nearly exclusively (95% of total N) as dissolved organic forms of nitrogen (DON), rather than as the inorganic forms N[O.sub.[3.sup.-]] (0.2% of total N) and N[H.sub.[4.sup.+]] (4.8% of total N). The strong dominance of organic over inorganic losses of N indicates that losses of N from old-growth forest ecosystems are not exclusively subject to traditional mechanisms of direct biotic control (i.e., mineralization supply or biotic uptake), but are also subject to indirect biotic control associated with the long-term accumulation, humification, and leaching of soil organic N during ecosystem succession. The view of unpolluted old-growth forests as "leaky" vs. "non-leaky" with respect to N depends on whether ecosystem budgets or models consider dissolved organic, as well as inorganic, forms of N. High- and mid-elevation forests in our study area showed the lowest efflux concentrations of N as N[O.sub.[3.sup.-]] reported from any old-growth temperate forest ecosystem (0.10 vs. 0.30 [µgram]/L, respectively). Comparisons against other old-growth forests, subject to varying levels of N deposition, indicated that levels of N[O.sub.[3.sup.-]] and the relative abundance of N[O.sub.[3.sup.-]] vs. N[H.sub.[4.sup.+]] in Northern Hemisphere forests may be strongly influenced by atmospheric N inputs. Our analyses suggest that patterns of N cycling, as well as evolutionary selective pressures on plant and microbial species, may have changed as a function of regional-scale increases in atmospheric N deposition to European and North American temperate forests. We hypothesize that, because they inherently are more poor than aggrading forests at retaining added nutrients, old-growth forest ecosystems are particularly sensitive indicators of N deposition.
Audience Academic
Author Armesto, Juan J.
Johnson, Arthur H.
Hedin, Lars O.
Author_xml – sequence: 1
  givenname: Lars O.
  surname: Hedin
  fullname: Hedin, Lars O.
– sequence: 2
  givenname: Juan J.
  surname: Armesto
  fullname: Armesto, Juan J.
– sequence: 3
  givenname: Arthur H.
  surname: Johnson
  fullname: Johnson, Arthur H.
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Copyright Copyright 1995 The Ecological Society of America
1995 by the Ecological Society of America
1995 INIST-CNRS
COPYRIGHT 1995 Ecological Society of America
Copyright Ecological Society of America Mar 1995
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– notice: 1995 by the Ecological Society of America
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– notice: COPYRIGHT 1995 Ecological Society of America
– notice: Copyright Ecological Society of America Mar 1995
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Keywords Nutrient
Air pollution
Biogeochemical cycle
Primary forest
Ecosystem
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PublicationTitle Ecology (Durham)
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Snippet Atmospheric pollution and other human activities have altered natural element cycles over large regions of the world. Much current understanding of nutrient...
The patterns of hydrologic nutrient loss from old-growth temperate forest ecosystems in southern Chile based on chemical analyses of small streams draining 31...
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SubjectTerms aerosols
air pollution
ammonium compounds
Animal and plant ecology
Animal, plant and microbial ecology
anthropogenic activities
Atmospherics
biocenosis
Biogeochemistry
Biological and medical sciences
Biology
Biotic communities
chemical analysis
Chile
Coastal forests
Coniferous forests
ecological succession
Ecosystems
Forest ecology
Forest ecosystems
Forest hydrology
Forest soils
Forests
Forests and forestry
Fundamental and applied biological sciences. Psychology
Geology
humification
hydrology
leaching
mineralization
nitrates
nitrogen
nitrogen content
Nutrient loss
nutrients
Old growth forests
prediction
soil organic nitrogen
streams
Synecology
Temperate forests
temperate zones
Terrestrial ecosystems
Watersheds
Title Patterns of Nutrient Loss from Unpolluted, Old-Growth Temperate Forests: Evaluation of Biogeochemical Theory
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