Deforestation of watersheds of Panama: nutrient retention and export to streams

A series of eight watersheds on the Pacific coast of Panama where conversion of mature lowland wet forest to pastures by artisanal burning provided watershed-scale experimental units with a wide range of forest cover (23, 29, 47, 56, 66, 73, 73, 91, and 92 %). We used these watersheds as a landscape-scale experiment to assess effects of degree of deforestation on within-watershed retention and hydrological export of atmospheric inputs of nutrients. Retention was estimated by comparing rainfall nutrient concentrations (volume-weighted to allow for evapotranspiration) to concentrations in freshwater reaches of receiving streams. Retention of rain-derived nutrients in these Panama watersheds averaged 77, 85, 80, and 62 % for nitrate, ammonium, dissolved organic N, and phosphate, respectively. Retention of rain-derived inorganic nitrogen, however, depended on watershed cover: retention of nitrate and ammonium in pasture-dominated watersheds was 95 and 98 %, while fully forested watersheds retained 65 and 80 % of atmospheric nitrate and ammonium inputs. Watershed forest cover did not affect retention of dissolved organic nitrogen and phosphate. Exports from more forested watersheds yielded DIN/P near 16, while pasture-dominated watersheds exported N/P near 2. The differences in magnitude of exports and ratios suggest that deforestation in these Panamanian forests results in exports that affect growth of plants and algae in the receiving stream and estuarine ecosystems. Watershed retention of dissolved inorganic nitrogen calculated from wet plus dry atmospheric deposition varied from 90 % in pasture- to 65 % in forest-dominated watersheds, respectively. Discharges of DIN to receiving waters from the watersheds therefore rose from 10 % of atmospheric inputs for pasture-dominated watersheds, to about 35 % of atmospheric inputs for fully forested watersheds. These results from watersheds with no agriculture or urbanization, but different conversion of forest to pasture by burning, show significant, deforestation-dependent retention within tropical watersheds, but also ecologically significant, and deforestation-dependent, exports that are biologically significant because of the paucity of nutrients in receiving tropical stream and coastal waters.     

高山森林溪流冬季不同时期凋落物分解中水溶性氮和磷的动态特征

为了解高山森林溪流凋落物冬季分解过程中水溶性氮和磷的变化过程,采用凋落叶分解袋法,以川西高山森林典型乔木(四川红杉、方枝柏)和灌木(高山杜鹃、康定柳)凋落叶为研究对象,研究冬季不同时期(冻结初期、冻结期、融化期)溪流、河流、河岸带以及林下凋落叶水溶性氮和磷的动态特征.结果表明:经过一个冬季的分解,4种凋落物在不同生境下各时期的水溶性氮含量无显著变化.水溶性磷含量除林下外在其他生境均显著降低,表现出河流＜溪流＜河岸带＜林下的规律.高山森林凋落叶分解过程中水溶性磷含量与平均温度、正积温、负积温和流速呈显著负相关,水溶性氮含量与正积温呈显著正相关,物种显著影响凋落物分解过程中水溶性氮和磷的含量.高山森林凋落物冬季分解过程中水溶性磷更易随河流和溪流等水体的流动而流失,而水溶性氮受冬季水环境的影响相对较小.     

Carbon and nitrogen stoichiometry and nitrogen cycling rates in streams

Stoichiometric analyses can be used to investigate the linkages between N and C cycles and how these linkages influence biogeochemistry at many scales, from components of individual ecosystems up to the biosphere. N-specific NH₄⁺ uptake rates were measured in eight streams using short-term ¹⁵N tracer additions, and C to N ratios (C:N) were determined from living and non-living organic matter collected from ten streams. These data were also compared to previously published data compiled from studies of lakes, ponds, wetlands, forests, and tundra. There was a significant negative relationship between C:N and N-specific uptake rate; C:N could account for 41% of the variance in N-specific uptake rate across all streams, and the relationship held in five of eight streams. Most of the variation in N-specific uptake rate was contributed by detrital and primary producer compartments with large values of C:N and small values for N-specific uptake rate. In streams, particulate materials are not as likely to move downstream as dissolved N, so if N is cycling in a particulate compartment, N retention is likely to be greater. Together, these data suggest that N retention may depend in part on C:N of living and non-living organic matter in streams. Factors that alter C:N of stream ecosystem compartments, such as removal of riparian vegetation or N fertilization, may influence the amount of retention attributed to these ecosystem compartments by causing shifts in stoichiometry. Our analysis suggests that C:N of ecosystem compartments can be used to link N-cycling models across streams.     

Biogeochemical responses of two forest streams to a 2-month calcium addition

1. Calcium (Ca) has been lost from forest soils at the Hubbard Brook Experimental Forest (HBEF) because of decreased atmospheric input of Ca and high input of acid anions. Through time, this Ca loss has led to low streamwater Ca concentration and this change may affect stream ecosystem processes.
2. To test both the biogeochemical response of streams to increased calcium concentration and the role of streams in retaining calcium lost from soils, we added c. 120 μeq Ca L^?1 as CaCl₂ to two second‐order streams at HBEF for 2 months. One stream (buffered) also received an equivalent amount of NaHCO₃ to simulate the increase in pH and alkalinity if Ca were added with associated HCO₃^? ion. The other stream (unbuffered) received only CaCl₂. We collected water samples along a transect above and below the addition site at 11 dates: two before, seven during, and two after the addition.
3. Increase in pH in the buffered stream ranged from 5.6 to about 7.0 in the treated section. There was a net uptake of Ca on all sampling dates during the addition and these uptake rates were positively related to pH. Between 10 and 50% of the added Ca was taken up during the release in the 80‐m study reach. In the unbuffered stream, there was net uptake of Ca on only two dates, suggesting lower Ca uptake.
4. Water samples collected after the addition was stopped showed that a small fraction of the added Ca desorbed from sediments; the remainder was apparently in longer‐term storage in the sediments. No Ca desorbed from the stream sediments in the unbuffered stream, showing that sorption/desorption may be controlled by a pH‐induced increase in the number of exchange sites.
5. These streams appeared to be a significant sink for Ca over a 2‐month time scale, and thus, change in streamwater Ca during a year may be due to processing of Ca within the stream channel, as well as to changes in inputs from the catchment.     

Computation and visualization of regional-scale forest disturbance and associated dissolved nitrogen export from Shenandoah National Park,Virginia

Long-term watershed research conducted in Shenandoah National Park (SNP) in Virginia and elsewhere in the eastern U.S. indicates that annual export of dissolved nitrogen (N) from gaged forested watersheds to surface waters increases dramatically in response to vegetation disturbances. Dissolved N leakage is a common, well-documented response of small forested watersheds to logging in the larger region, while recent defoliation outbreaks of the gypsy moth ( Lymantria dispar) larva in the deciduous forests of SNP have been shown to generate similar biogeochemical responses. A recent modeling analysis further suggests that a parsimonious, empirical, unit N export response function (UNERF) model can explain large percentages of the temporal variation in annual N export from a group of small gaged forested watersheds in the years following disturbance. The empirical UNERF modeling approach is completely analogous to the unit hydrograph technique for describing storm runoff, with the model representing annual N export as a linear deterministic process both in space and in time. The purposes of this analysis are to (1) test the applicability of the UNERF model using quarterly streamwater nitrate data from a group of ungaged watersheds in SNP; (2) demonstrate a park-wide application of a regional UNERF model that references the geographic distributions of bedrock geology and the timing and extent of gypsy moth defoliation over the entire SNP area; and (3) visualize the temporal and spatial patterns in vegetation disturbance and annual dissolved N export through the use of computer animation software. During water year 1992, the year of peak defoliation, our modeling study suggests that park-wide export had transiently increased by 1700% from a baseline rate of about 0.1 kg/ha/year. SNP forests appear to be characteristic of other N-limited second-growth forests in the eastern U.S. that leak little N under undisturbed conditions, despite receiving relatively large inputs of N from atmospheric deposition sources. Vegetation disturbances can apparently cause major changes in N input-output balances with potentially important ramifications for low-order forest streams and downstream receiving waters.     

Remarks on the effects of nitrogen deposition to forest ecosystems

The effects of increased deposition of nitrogen compounds on forest sites are discussed based on literature data and own results from both earlier fertilization experiments and the ARINUS study area in the Black Forest. The influence on mycorrhiza is stated suppressive as well as stimulating so that no general conclusion can be drawn. The nitrogen nutrition status of coniferous forests is still sub-optimal over wide areas with a yearly deposition of 10–20 kg N ha^-1. Under considerable higher input rates the insufficient supply of other nutrients and imbalances in the nutritional status of trees are possible. When discussing nitrogen saturation of ecosystems, the nitrogen storage capacity of soils has to be considered as a decisive factor which varies from site to site. Any actual input/output balance is strongly influenced by the internal turnover processes resulting from former land use.     

Trophic structure of macro- and meso-invertebrates in Japanese coniferous forest: Carbon and nitrogen stable isotopes analyses

Few studies have been published on the feeding ecology of Japanese soil fauna based on stable isotope analysis. Therefore, the present work aims to use this technique for studying the trophic structure of Japanese soil fauna at two coniferous forests. Significant differences were observed between investigated sites (Arahama and Gamo) in genus richness and abundance, while for Shannon diversity indexes the difference was non-significant. The isotopic signatures (δ¹³C and δ¹⁵N) of the invertebrates collected at Arahama ranged from 0.3 to 6.3‰ for δ¹⁵N and from −27.3 to −23.3‰ for δ¹³C. At Gamo, invertebrates δ¹³C values ranged from −26.1 to −23.5‰ and δ¹⁵N values ranged from 1.6 to 6.8‰. At both sites, invertebrates formed two distinct groups on the basis of combined C and N stable isotope ratios. The locations of these groups related to δ¹³C values. The less enriched group (δ¹³C < −25‰) and the more enriched one (δ¹³C > −25‰). The range of δ¹⁵N for the present animals exceeded two trophic levels. While, the gradual ¹⁵N enrichment within the invertebrates species may indicate the dominance of omnivory in soil food webs. The differences between sites in δ¹⁵N confirm the importance of studying the trophic structure of soil fauna locally.     

Thermo-erosion gullies increase nitrogen available for hydrologic export

Formation of thermokarst features, ground subsidence caused by thaw of ice-rich permafrost, can result in increased export of inorganic nitrogen (N) from arctic tundra to downstream ecosystems. We compared physical characteristics, N pools, and rates of N transformations in soils collected from thermo-erosion gullies, intact water tracks (the typical precursor landform to thermo-erosion gullies), and undisturbed tundra to test potential mechanisms contributing to export of inorganic N. Subsidence exposes mineral soils, which tend to contain higher abundance of inorganic ions relative to surface soils, and may bring inorganic N into contact with flowing water. Alternatively, physical mixing may increase aeration and drainage of soils, which could promote N mineralization and nitrification while suppressing denitrification. Finally, some soil types are more prone to formation of thermokarst, and if these soils are relatively N-rich, thermokarst features may export more N than surrounding tundra. Inorganic N pools in thermo-erosion gullies were similar to the mean for all tundra types in this region, as well as to water tracks when integrated across two sampled depths. Thus, soils prone to thermo-erosion are not intrinsically N-rich, and increased N availability in thermokarst features is apparent only at sub-regional spatial scales. However, vertical profiles of N pools and transformation rates were homogenized within thermo-erosion gullies compared to adjacent intact tundra, indicating that physical mixing brings inorganic N to the surface, where it may be subject to hydrologic export. Increased inorganic N availability caused by formation of thermo-erosion gullies may have acute, localized consequences for aquatic ecosystems downstream of positions within drainage networks that are susceptible to thermo-erosion.     

Contribution of amino compounds to dissolved organic nitrogen in forest soils

Dissolved organic nitrogen (DON) may play an important role in plantnutrition and nitrogen fluxes in forest ecosystems. In spite of the apparentimportance of DON, there is a paucity of information concerning its chemicalcomposition. However, it is exactly this chemical characterization that isrequired to understand the importance of DON in ecosystem processes. Theprimaryobjective of this study was to characterize the distribution of free aminoacidsand hydrolyzable peptides/proteins in the DON fraction of Oa horizon leachatesalong an extreme edaphic gradient in northern California. Insitu soil solutions were extracted by centrifugation from Oahorizonscollected beneath Pinus muricata (Bishop pine) andCupressus pygmaea (pygmy cypress) on slightlyacidic/fertile and highly acidic/infertile sites. DON accounted for 77 to99% of the total dissolved nitrogen in Oa horizon leachates. Nitrogen infree amino acids and alkyl amines ranged from 0.04–0.07 mgN/L on the low fertility site to 0.45–0.49 mg N/L onthe high fertility site, and accounted for 1.5 to 10.6% of the DON fraction.Serine, glutamic acid, leucine, ornithine, alanine, aspartic acid andmethylamine were generally the most abundant free amino compounds. Combinedamino acids released by acid hydrolysis accounted for 48 to 74% of theDON, suggesting that proteins and peptides were the main contributor to DON inOa horizon leachates. Together, nitrogen from free andcombined amino compounds accounted for 59 to 78% of the DON. Most of theDON was found in the hydrophobic fraction, which suggests the presence ofprotein/peptide-polyphenol complexes or amino compounds associated withhumic substances. Because free and combined amino acids can be an importantnitrogen source for some plants, soil DON may play an important role in plantnutrition and ecosystem function.     

Anthropogenic nitrogen sources and relationships to riverine nitrogen export in the northeastern U.S.A.

Human activities have greatly altered the nitrogen (N) cycle, accelerating the rate of N fixation in landscapes and delivery of N to water bodies. To examine relationships between anthropogenic N inputs and riverine N export, we constructed budgets describing N inputs and losses for 16 catchments, which encompass a range of climatic variability and are major drainages to the coast of the North Atlantic Ocean along a latitudinal profile from Maine to Virginia. Using data from the early 1990's, we quantified inputs of N to each catchment from atmospheric deposition, application of nitrogenous fertilizers, biological nitrogen fixation, and import of N in agricultural products (food and feed). We compared these inputs with N losses from the system in riverine export.The importance of the relative sources varies widely by catchment and is related to land use. Net atmospheric deposition was the largest N source (>60%) to the forested basins of northern New England (e.g. Penobscot and Kennebec); net import of N in food was the largest source of N to the more populated regions of southern New England (e.g. Charles & Blackstone); and agricultural inputs were the dominant N sources in the Mid-Atlantic region (e.g. Schuylkill & Potomac). Over the combined area of the catchments, net atmospheric deposition was the largest single source input (31%), followed by net imports of N in food and feed (25%), fixation in agricultural lands (24%), fertilizer use (15%), and fixation in forests (5%). The combined effect of fertilizer use, fixation in crop lands, and animal feed imports makes agriculture the largest overall source of N. Riverine export of N is well correlated with N inputs, but it accounts for only a fraction (25%) of the total N inputs. This work provides an understanding of the sources of N in landscapes, and highlights how human activities impact N cycling in the northeast region.     

Fine root turnover in forest ecosystems in relation to quantity and form of nitrogen availability: a comparison of two methods

Summary Two methods of estimating fine root production and turnover are compared for 13 forest ecosystems exhibiting a wide range in form (NH₄ ⁺ vs. NO₃ ^-) and quantity of available nitrogen. The two methods are by comparison of seasonal maxima and minima in biomess and by nitrogen budgeting. Both methods give similar results for stands with low rates of nitrification. The budgeting method predicts higher fine root turnover and productivity than the max-min method for systems with significant rates of nitrification.     

Algal nitrogen fixation in Californian streams: seasonal cycles

Using the acetylene reduction technique, nitrogen fixation was measured in Rocky Creek, a small seasonally dry Californian stream. In the 3 years since 1970 nitrogen fixation varied seasonally and spatially, being highest in the early stages of colony growth in shallow, clear regions where there was little shade. The annual rate of N₂-fixation was similar to that found recently for Arctic tundra, but was greater than rates for Antarctic rivulets and less than rates for temperate rocky shores. A hetero-cystous Nostoc appeared to be the sole organism responsible for this fixation. Nostoc occurred in variously sized gelatinous clumps on the stable boulders on which the co-dominant alga, Ulothrix zonata, also grew. Maximum nitrogenase activity yielded 70 nmoles C₂H₄ mg^-1 drywt day^-1 and 123 nmoles C₂H₄cm^-2 day^-1. Estimated annual amounts of nitrogen fixed in 1971 ranged from 42 in shade to 360 mg N fixed m^-2y^-1 in the most favourable areas. Nostoc biomass reached 33 g dry wt m^-2 and peeled off each year in June-July. Because Nostoc was confined to stable rocks the biomass per unit area of the stream as a whole was much less, ranging from 0.054 to 1.26 g dry wt m^-2 in the most favourable site. Nostoc was common throughout the length of Rocky Creek and also plentiful in eight out of ten adjacent streams but not in the main river (Eel River South Fork), probably due to high turbulence and turbidity. N₂-fixation in these streams makes a significant, but probably small contribution to the nitrogen income of the nitrogen-deficient Eel River system. Nostoc colony establishment appears to be controlled by a combination of reduced turbulence and firmer attachment of the colonies to the substrate. Nostoc colony disappearance in June is probably due to nutrient depletion.     

Immediate and long-term nitrogen oxide emissions from tropical forest soils exposed to elevated nitrogen input

Tropical nitrogen (N) deposition is projected to increase substantially within the coming decades. Increases in soil emissions of the climate‐relevant trace gases NO and N₂O are expected, but few studies address this possibility. We used N addition experiments to achieve N‐enriched conditions in contrasting montane and lowland forests and assessed changes in the timing and magnitude of soil N‐oxide emissions. We evaluated transitory effects, which occurred immediately after N addition, and long‐term effects measured at least 6 weeks after N addition. In the montane forest where stem growth was N limited, the first‐time N additions caused rapid increases in soil N‐oxide emissions. During the first 2 years of N addition, annual N‐oxide emissions were five times (transitory effect) and two times (long‐term effect) larger than controls. This contradicts the current assumption that N‐limited tropical montane forests will respond to N additions with only small and delayed increases in soil N‐oxide emissions. We attribute this fast and large response of soil N‐oxide emissions to the presence of an organic layer (a characteristic feature of this forest type) in which nitrification increased substantially following N addition. In the lowland forest where stem growth was neither N nor phosphorus (P) limited, the first‐time N additions caused only gradual and minimal increases in soil N‐oxide emissions. These first N additions were completed at the beginning of the wet season, and low soil water content may have limited nitrification. In contrast, the 9‐ and 10‐year N‐addition plots displayed instantaneous and large soil N‐oxide emissions. Annual N‐oxide emissions under chronic N addition were seven times (transitory effect) and four times (long‐term effect) larger than controls. Seasonal changes in soil water content also caused seasonal changes in soil N‐oxide emissions from the 9‐ and 10‐year N‐addition plots. This suggests that climate change scenarios, where rainfall quantity and seasonality change, will alter the relative importance of soil NO and N₂O emissions from tropical forests exposed to elevated N deposition.     

Macroinvertebrate community response to natural and forest harvest gradients in western Oregon headwater streams

1. To examine the effects of forest harvest practices on headwater stream macroinvertebrates, we compiled a 167 site database with macroinvertebrate, fish, physical habitat and catchment land cover data from the three forested ecoregions in western Oregon. For our analysis, headwater streams were defined by catchment areas <10 km² and perennial water during summer low flows. Almost all sites in the database were selected using a randomised survey design, constituting a representative sample of headwater streams in these ecoregions. 2. Macroinvertebrate taxonomic and functional feeding group composition were very similar among the three ecoregions in the study area (Coast Range, Cascades and Klamath Mountains). On average, 55% of the individuals at each site were in the orders Ephemeroptera, Plecoptera or Trichoptera. Dipteran taxa (mostly chironomids) accounted for another 34%. At almost all sites, non‐insects made up <10% of the macroinvertebrate assemblage. Almost half (49%) of the assemblages were collectors; remaining individuals were about evenly divided among scrapers, shredders and predators. 3. There were 189 different macroinvertebrate taxa at the 167 sites with richness at individual sites ranging from 7 to 71 taxa. Ordination by non‐metric multidimensional scaling revealed a strong association between % Ephemeroptera, especially Baetis, and site scores along the first axis. This axis was also strongly related to % coarse substratum and fast water habitat. The second axis was strongly related to % intolerant individuals, site slope and altitude. No strong relationships were evident between any ordination axis and either logging activity, presence/absence of fish, catchment size or ecoregion. 4. Based on macroinvertebrate index of biotic integrity (IBI) scores, 62% of the sites had no impairment, 31% of the sites had slight impairment and only 6% of the sites had moderate or severe impairment. IBI scores were not strongly related to forest harvest history. All four severely impaired sites and five of the seven sites with moderate impairment were lower altitude, shallower slope stream reaches located in the Coast Range with evidence of agricultural activity in their catchment or riparian zone. % sand + fine substratum was the environmental variable most strongly related to macroinvertebrate IBI.     

Retention of nitrogen in small streams artificially polluted with nitrate

A simple method was developed to test hypotheses on nitrogen retention in first-order streams in an agricultural region near Oslo, SE Norway. A gravity-operated system added a nitrate solution to the streams continuously at a constant rate. Water samples were collected at fixed intervals downstream to follow the rate of decline in streamwater nitrate. Repeated sampling allowed calculation of regression lines from experiments with different levels of additions of nitrate.The experiments showed that removal of nitrate generally increased with higher initial nitrate concentration, regardless of temperature (range 8–16 °C). Higher nitrate removal rates were found in a stream polluted by easily degradable organic matter than in a similar stream fed by groundwater.Experiments conducted in indoor channels lined with a layer of stream sediment gave reproducible, exponential rates of nitrate decrease in the recirculated water.The results are discussed in the framework of first-order streams as protective ecotones between agricultural areas and higher-order parts of the watersheds.     

Impact of winter climate change on nitrogen biogeochemistry in forest ecosystems: A synthesis from Japanese case studies

Nitrogen (N) is a critical ecological and environmental indicator under changing environments. The impact of winter climate change on N biogeochemical processes in forest ecosystems has gained increasing recognition. Decreasing snowfall has caused a decrease in the heat insulation properties of the snowpack, resulting in an increase in the frequency and magnitude of freezing and thawing cycles in surface soil, where biological processes are most active. Here I synthesize recent research findings from integrated field observations and experiments conducted in northern Japan and compare these results with previous research outcomes from other regions to identify current research gaps and develop the next research agenda to further advance our understanding of this complex problem. Japanese case studies indicated that net ammonium production (ammonification) was mostly dominant in terms of available soil N fertility in cold environments and was sensitive to the increase in soil freezing and thawing cycles because of the decreased snowpack. On the other hands, nitrate dynamics were more stable or conservative than those of ammonium. The soil characteristics (i.e., N pool and microbial activities) were significant explanatory factors of the responses of soil N dynamics and N leakage among different soils to increased freezing–thawing cycles at watershed and national scale. This synthesis indicates that winter climate change had significant impacts on soil N biogeochemistry (such as soil N pool size and microbial N transformation) during the winter and snowmelt season and also during the following growing season. Several research gaps and possible research topics (path dependency and soil microbial community composition) are also presented by synthesizing the current research findings. Further field experiments and observations quantifying the pools and fluxes of inorganic N with modeling analysis under freeze–thaw environments would contribute to increase the understandings of N transformation processes under winter climate change.