On linking multiyear biometric measurements of tree growth with eddy covariance-based net ecosystem production

Annual measurements of the diameter growth and litter fall of trees began in 1998 using a 1.0 ha permanent plot beneath a flux tower at the Takayama flux site, central Japan. This opened up an opportunity for studies that compare the interannual variability in tree growth with eddy covariance-based net ecosystem production (NEP). A possible link between multiyear biometric-based net primary production (NPP) and eddy covariance-based NEP was investigated to determine the contribution of autotrophic production and heterotrophic respiration (HR) to the interannual variability of NEP in deciduous forest ecosystems. We also defined the NEP^* as the measurable organic matter stored in an ecosystem during the interval in which soil respiration (SR) measurements were taken. The difference of biometric-based NEP^* from eddy covariance-based NEP within a given year varied between 55% and 105%. Woody tissue NPP (stems and coarse roots) varied markedly from 0.88 to 1.96 Mg C ha⁻¹ yr⁻¹ during the 8-year study period (1999–2006). Annual woody tissue NPP was positively correlated with eddy covariance-based NEP ( r ²=0.52, P <0.05). However, neither foliage NPP ( r ²=0.03) nor HR ( r ²=0.06) were correlated with eddy covariance-based NEP. Therefore, it was hypothesized that interannual variability in the ecosystem carbon exchange was directly responsible for much of the interannual variation in autotrophic production, especially carbon accumulation in the woody components of the ecosystem. Moreover, similar interannual variations of biometric-based NEP^* and eddy covariance-based NEP with small variations in SR and foliage NPP suggest a constant net accumulation of carbon in nonliving pools at the Takayama site.     

Carbon mitigation by the energy crop, Miscanthus

Biomass crops mitigate carbon emissions by both fossil fuel substitution and sequestration of carbon in the soil. We grew Miscanthus x giganteus for 16 years at a site in southern Ireland to (i) compare methods of propagation, (ii) compare response to fertilizer application and quantify nutrient offtakes, (iii) measure long-term annual biomass yields, (iv) estimate carbon sequestration to the soil and (v) quantify the carbon mitigation by the crop. There was no significant difference in the yield between plants established from rhizome cuttings or by micro-propagation. Annual off-takes of N and P were easily met by soil reserves, but soil K reserves were low in unfertilized plots. Potassium deficiency was associated with lower harvestable yield. Yields increased for 5 years following establishment but after 10 years showed some decline which could not be accounted for by the climate driven growth model MISCANMOD. Measured yields were normalized to estimate both autumn (at first frost) and spring harvests (15 March of the subsequent year). Average autumn and spring yields over the 15 harvest years were 13.4±1.1 and 9.0±0.7 t DW ha⁻¹ yr⁻¹ respectively. Below ground biomass in February 2002 was 20.6±4.6 t DW ha⁻¹. Miscanthus derived soil organic carbon sequestration detected by a change in ¹³C signal was 8.9±2.4 t C ha⁻¹ over 15 years. We estimate total carbon mitigation by this crop over 15 years ranged from 5.2 to 7.2 t C ha⁻¹ yr⁻¹ depending on the harvest time.     

Terrestrial export of highly bioavailable carbon from small boreal catchments in spring floods

1. We assessed the terrestrial export of organic carbon, which effectively supported aquatic bacterial production (BP), from small boreal catchments during spring flood. We analysed stream runoff from nine small catchments with different proportions of peat mires and coniferous forests by monitoring the dissolved organic carbon (DOC) flux in combination with conducting bacterial bioassays.
2. Multiple linear regression analysis showed that BP during 7-day-dark bioassays (BP₇; μg C L⁻¹day⁻¹) was explained by both the quantity and quality (low-molecular weight fractions) of the DOC. BP₇ can be used as a measure of export of terrestrial organic carbon that is highly bioavailable.
3. Total export of DOC during spring flood from the different catchments ranged from 20 to 27 kg ha⁻¹ and was negatively correlated to forest cover (%). However, the export of BP₇ carbon was positively correlated to forest cover and varied from about 0.1 kg ha⁻¹ in mire-dominated streams to about 0.2 kg ha⁻¹ in forest-dominated streams.
4. The high bioavailability of forest carbon suggests that forests are the main contributors of BP-supporting carbon in boreal streams although mires have higher area-specific export of DOC.     

Change in CO2 balance under a series of forestry activities in a cool-temperate mixed forest with dense undergrowth

To evaluate the effects on CO₂ exchange of clearcutting a mixed forest and replacing it with a plantation, 4.5 years of continuous eddy covariance measurements of CO₂ fluxes and soil respiration measurements were conducted in a conifer-broadleaf mixed forest in Hokkaido, Japan. The mixed forest was a weak carbon sink (net ecosystem exchange, −44 g C m⁻² yr⁻¹), and it became a large carbon source (569 g C m⁻² yr⁻¹) after clearcutting. However, the large emission in the harvest year rapidly decreased in the following 2 years (495 and 153 g C m⁻² yr⁻¹, respectively) as the gross primary production (GPP) increased, while the total ecosystem respiration (RE) remained relatively stable. The rapid increase in GPP was attributed to an increase in biomass and photosynthetic activity of Sasa dwarf bamboo, an understory species. Soil respiration increased in the 3 years following clearcutting, in the first year mainly owing to the change in the gap ratio of the forest, and in the following years because of increased root respiration by the bamboo. The ratio of soil respiration to RE increased from 44% in the forest to nearly 100% after clearcutting, and aboveground parts of the vegetation contributed little to the RE although the respiration chamber measurements showed heterogeneous soil condition after clearcutting.     

Changes in topsoil carbon stock in the Tibetan grasslands between the 1980s and 2004

Climate warming is likely inducing carbon loss from soils of northern ecosystems, but little evidence comes from large-scale observations. Here we used data from a repeated soil survey and remote sensing vegetation index to explore changes in soil organic carbon (SOC) stock on the Tibetan Plateau during the past two decades. Our results showed that SOC stock in the top 30 cm depth in alpine grasslands on the plateau amounted to 4.4 Pg C (1 Pg=10¹⁵ g), with an overall average of 3.9 kg C m⁻². SOC changes during 1980s–2004 were estimated at −0.6 g C m⁻² yr⁻¹, ranging from −36.5 to 35.8 g C m⁻² yr⁻¹ at 95% confidence, indicating that SOC stock in the Tibetan alpine grasslands remained relatively stable over the sampling periods. Our findings are nonconsistent with previous reports of loss of soil C in grassland ecosystems due to the accelerated decomposition with warming. In the case of the alpine grasslands on the Tibetan Plateau studied here, we speculate that increased rates of decomposition as soils warmed during the last two decades may have been compensated by increased soil C inputs due to increased grass productivity. These results suggest that soil C stock in terrestrial ecosystems may respond differently to climate change depending on ecosystem type, regional climate pattern, and intensity of human disturbance.     

Simulated chronic nitrogen deposition increases carbon storage in Northern Temperate forests

High levels of atmospheric nitrogen (N) deposition in Europe and North America were maintained throughout the 1990s, and global N deposition is expected to increase by a factor of 2.5 over the next century. Available soil N limits primary production in many terrestrial ecosystems, and some computer simulation models have predicted that increasing atmospheric N deposition may result in greater terrestrial carbon (C) storage in woody biomass. However, empirical evidence demonstrating widespread increases in woody biomass C storage due to atmospheric N deposition is uncommon. Increased C storage in soil organic matter due to chronic N inputs has rarely been reported and is often not considered in computer simulation models of N deposition effects. Since 1994, we have experimentally simulated chronic N deposition by adding 3 g N m⁻² yr⁻¹ to four different northern hardwood forests, which span a 500 km geographic gradient in Michigan. Each year we measured tree growth. In 2004, we also examined soil C content to a depth of 70 cm. When we compared the control treatment with the NO₃⁻ deposition treatment after a decade of experimentation, ecosystem C storage had significantly increased in both woody biomass (500 g C m⁻²) and surface soil (0–10 cm) organic matter (690 g C m⁻²). The increase in surface soil C storage was apparently driven by altered rates of organic matter decomposition, rather than an increase in detrital inputs to soil. Our results, for study locations stretching across hundreds of kilometers, support the hypothesis that chronic N deposition may increase C storage in northern forests, potentially contributing to a sink for anthropogenic CO₂ in the northern Hemisphere.     

Climate-induced changes in high elevation stream nitrate dynamics

Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991–1999. Mean annual N export increased by 28% from 2.03 kg N ha⁻¹ yr⁻¹ before 2000 to 2.84 kg N ha⁻¹ yr⁻¹ in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha⁻¹ yr⁻¹) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12 °C yr⁻¹ in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000–2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat.     

Carbon pool densities and a first estimate of the total carbon pool in the Mongolian forest‐steppe

The boreal forest biome represents one of the most important terrestrial carbon stores, which gave reason to intensive research on carbon stock densities. However, such an analysis does not yet exist for the southernmost Eurosiberian boreal forests in Inner Asia. Most of these forests are located in the Mongolian forest‐steppe, which is largely dominated by Larix sibirica. We quantified the carbon stock density and total carbon pool of Mongolia's boreal forests and adjacent grasslands and draw conclusions on possible future change. Mean aboveground carbon stock density in the interior of L. sibirica forests was 66 Mg C ha^?1, which is in the upper range of values reported from boreal forests and probably due to the comparably long growing season. The density of soil organic carbon (SOC, 108 Mg C ha^?1) and total belowground carbon density (149 Mg C ha^?1) are at the lower end of the range known from boreal forests, which might be the result of higher soil temperatures and a thinner permafrost layer than in the central and northern boreal forest belt. Land use effects are especially relevant at forest edges, where mean carbon stock density was 188 Mg C ha^?1, compared with 215 Mg C ha^?1 in the forest interior. Carbon stock density in grasslands was 144 Mg C ha^?1. Analysis of satellite imagery of the highly fragmented forest area in the forest‐steppe zone showed that Mongolia's total boreal forest area is currently 73 818 km², and 22% of this area refers to forest edges (defined as the first 30 m from the edge). The total forest carbon pool of Mongolia was estimated at ~ 1.5?1.7 Pg C, a value which is likely to decrease in future with increasing deforestation and fire frequency, and global warming.     

Storms can cause Europe-wide reduction in forest carbon sink

Disturbance of ecosystems is a major factor in regional carbon budgets, and it is believed to be partly responsible for the large inter-annual variability of the terrestrial part of the carbon balance. Forest fires have so far been considered as the most important disturbance but also other forms of disturbance such as insect outbreaks or wind-throw might contribute significantly to the largely unexplained inter-annual variability, at least in specific regions. The effect of wind-throw has not yet been estimated because of lack of data on how carbon fluxes are affected. The Gudrun storm, which hit Sweden in January 2005, resulted in ca. 66 million m³ of wind-thrown stem wood on an area of ca. 272 000 ha. Using a model (BIOME-BGC) calibrated to CO₂ flux measurements at two sites, the annual net ecosystem productivity during the first year after the storm was estimated to be in the range −897 to −1259 g C m⁻² yr⁻¹. This is a much higher loss compared with harvested (clear-cut) forests in Europe, which ranged between ca. −420 and −100 g m⁻² yr⁻¹. The reduction in the carbon sink scaled to the whole wind-thrown area was estimated at ca. 3 million tons C during the first year. By historical data on wind-throw in Europe combined with modelling, we estimated that the large Lothar storm in 1999 reduced the European carbon balance by ca. 16 million tons C, this is ca. 30% of the net biome production in Europe. We conclude that the impact of increased forest damage by more frequent storms in future climate change scenarios must be considered and that intermittent large wind-throw events may explain a part of the large inter-annual variability in the terrestrial carbon sink.     

Holocene carbon burial by lakes in SW Greenland

The role of the Arctic in future global change processes is predicted to be important because of the large carbon (C) stocks contained in frozen soils and peatlands. Lakes are an important component of arctic landscapes although their role in storing C is not well prescribed. The area around Kangerlussuaq, SW Greenland (66–68°N, 49–54°W) has extremely high lake density, with ∼20 000 lakes that cover about 14% of the land area. C accumulation rates and standing stock (kg C m⁻²), representing late- to mid-Holocene C burial, were calculated from AMS ¹⁴C-dated sediment cores from 11 lakes. Lake ages range from ∼10 000 cal yr  bp to ∼5400 cal yr  bp , and reflect the withdrawal of the ice sheet from west to east. Total standing stock of C accumulated in the studied lakes for the last ∼8000 years ranged from 28 to 71 kg C m⁻², (mean: ∼42 kg C m⁻²). These standing stock determinations yield organic C accumulation rates of 3.5–11.5 g C m⁻² yr⁻¹ (mean: ∼6 g C m⁻² yr⁻¹) for the last 4500 years. Mean C accumulation rates are not different for the periods 8–4.5 and 4.5–0 ka, despite cooling trends associated with the neoglacial period after 4.5 ka. We used the mean C standing stock to estimate the total C pool in small lakes (<100 ha) of the Kangerlussuaq region to be ∼4.9 × 10¹³ g C. This C stock is about half of that estimated for the soil pool in this region (but in 5% of the land area) and indicates the importance of incorporating lakes into models of regional C balance at high latitudes.     

The global relationship between forest productivity and biomass

Aim  We aim to determine the empirical relationship between above-ground forest productivity and biomass. There are theoretical reasons to assume a relationship between forest structure and function, as both may be influenced by similar ecological factors such as moisture supply. Also, dynamic global vegetation model simulations imply that any increase in forest productivity driven by climate change will result in increases in biomass and therefore carbon storage. However, few studies have explored the strength and form of the relationship between forest productivity and biomass, whether in space or time.
Location Global scale.
Methods  We collated a large data set of above-ground biomass (AGB) and above-ground net primary productivity (ANPP) and tested the extent to which spatial variation in forest biomass across the Earth can be predicted from forest productivity.
Results  The global ANPP–AGB relationship differs fundamentally from the strongly positive, linear relationship reported in earlier analyses, which mostly lacked tropical sites. AGB begins to peak at c . 15–20 Mg ha⁻¹ year⁻¹ ANPP, plateaus at ANPP > 20–25 Mg ha⁻¹ year⁻¹, and may actually decline at higher levels of production.
Main conclusions  High turnover rates in high-productivity forests may limit AGB by promoting the dominance of species with a low wood density. Predicted increases in ANPP will not necessarily favour increases in forest carbon storage, especially if changes in productivity are accompanied by compositional shifts.     

Changes in soil organic carbon under biofuel crops

One potentially significant impact of growing biofuel crops will be the sequestration or release of carbon (C) in soil. Soil organic carbon (SOC) represents an important C sink in the lifecycle C balances of biofuels and strongly influences soil quality. We assembled and analyzed published estimates of SOC change following conversion of natural or agricultural land to biofuel crops of corn with residue harvest, sugarcane, Miscanthus x giganteus , switchgrass, or restored prairie. We estimated SOC losses associated with land conversion and rates of change in SOC over time by regressing net change in SOC relative to a control against age since establishment year. Conversion of uncultivated land to biofuel agriculture resulted in significant SOC losses – an effect that was most pronounced when native land was converted to sugarcane agriculture. Corn residue harvest (at 25–100% removal) consistently resulted in SOC losses averaging 3–8 Mg ha⁻¹ in the top 30 cm, whereas SOC accumulated under all four perennial grasses, with SOC accumulation rates averaging <1 Mg ha⁻¹ yr⁻¹ in the top 30 cm. More intensive harvests led to decreased C gains or increased C losses – an effect that was particularly clear for residue harvest in corn. Direct or indirect conversion of previously uncultivated land for biofuel agriculture will result in SOC losses that counteract the benefits of fossil fuel displacement. Additionally, SOC losses under corn residue harvest imply that its potential to offset C emissions may be overestimated, whereas SOC sequestration under perennial grasses represents an additional benefit that has rarely been accounted for in life cycle analyses of biofuels.     

Quantifying above‐ and belowground biomass carbon loss with forest conversion in tropical lowlands of Sumatra (Indonesia)

Natural forests in South‐East Asia have been extensively converted into other land‐use systems in the past decades and still show high deforestation rates. Historically, lowland forests have been converted into rubber forests, but more recently, the dominant conversion is into oil palm plantations. While it is expected that the large‐scale conversion has strong effects on the carbon cycle, detailed studies quantifying carbon pools and total net primary production (NPP_total) in above‐ and belowground tree biomass in land‐use systems replacing rainforest (incl. oil palm plantations) are rare so far. We measured above‐ and belowground carbon pools in tree biomass together with NPP_total in natural old‐growth forests, ‘jungle rubber’ agroforests under natural tree cover, and rubber and oil palm monocultures in Sumatra. In total, 32 stands (eight plot replicates per land‐use system) were studied in two different regions. Total tree biomass in the natural forest (mean: 384 Mg ha^?1) was more than two times higher than in jungle rubber stands (147 Mg ha^?1) and >four times higher than in monoculture rubber and oil palm plantations (78 and 50 Mg ha^?1). NPP_total was higher in the natural forest (24 Mg ha^?1 yr^?1) than in the rubber systems (20 and 15 Mg ha^?1 yr^?1), but was highest in the oil palm system (33 Mg ha^?1 yr^?1) due to very high fruit production (15–20 Mg ha^?1 yr^?1). NPP_total was dominated in all systems by aboveground production, but belowground productivity was significantly higher in the natural forest and jungle rubber than in plantations. We conclude that conversion of natural lowland forest into different agricultural systems leads to a strong reduction not only in the biomass carbon pool (up to 166 Mg C ha^?1) but also in carbon sequestration as carbon residence time (i.e. biomass‐C:NPP‐C) was 3–10 times higher in the natural forest than in rubber and oil palm plantations.     

Artificial drainage and associated carbon fluxes (CO2/CH4) in a tundra ecosystem

Ecosystem flux measurements using the eddy covariance (EC) technique were undertaken in 4 subsequent years during summer for a total of 562 days in an arctic wet tundra ecosystem, located near Cherskii, Far-Eastern Federal District, Russia. Methane (CH₄) emissions were measured using permanent chambers. The experimental field is characterized by late thawing of permafrost soils in June and periodic spring floods. A stagnant water table below the grass canopy is fed by melting of the active layer of permafrost and by flood water. Following 3 years of EC measurements, the site was drained by building a 3 m wide drainage channel surrounding the EC tower to examine possible future effects of global change on the tundra tussock ecosystem. Cumulative summertime net carbon fluxes before experimental alteration were estimated to be about +15 g C m⁻² (i.e. an ecosystem C loss) and +8 g C m⁻² after draining the study site. When taking CH₄ as another important greenhouse gas into account and considering the global warming potential (GWP) of CH₄ vs. CO₂, the ecosystem had a positive GWP during all summers. However CH₄ emissions after drainage decreased significantly and therefore the carbon related greenhouse gas flux was much smaller than beforehand (475 ± 253 g C-CO₂-e m⁻² before drainage in 2003 vs. 23 ± 26 g C-CO₂-e m⁻² after drainage in 2005).     

Near-zero recent carbon accumulation in a bog with high nitrogen deposition in SW Sweden

We present data on the accumulation of carbon and nitrogen into an open oceanic ombrotrophic bog, SW Sweden, with high levels of anthropogenic nitrogen deposition. The aim was to investigate if this peatland currently acts as a sink for atmospheric carbon. Peat cores were sampled from the top peat layer in five different vegetation types. Small pines were used to date the cores. The cores bulk density and carbon and nitrogen content were determined. A vegetation-classified satellite image was used to estimate the areal extent of the vegetation types and to scale up these results to bog level. The rate of current carbon input into the upper oxic acrotelm was 290 g m⁻² yr⁻¹, and there were no significant differences in accumulation rates among the vegetation types. This organic matter input to the acrotelm was almost completely decomposed before it was deposited for storage in the deeper peat layers (the catotelm) and only a small fraction (≪1%) or 0.012 g m⁻² yr⁻¹ of the carbon would be left, assuming a residence time of 100 years in the acrotelm. Nitrogen accumulation rates differed between the vegetation classes, and the average input via primary production varied from 5.33 to 16.8 g m⁻² yr⁻¹. Current nitrogen input rates into the catotelm are much lower, 0–0.059 g m⁻² yr⁻¹, with the highest accumulation rates in lawn-dominated communities. We suggest that one of the main causes of the low carbon input rates is the high level of nitrogen deposition, which enhances decomposition and changes the vegetation from peat-forming Sphagnum -dominance to dominance by dwarf shrubs and graminoids.     

Convergence in the relationship of CO2 and N2O exchanges between soil and atmosphere within terrestrial ecosystems

Ecosystem CO₂ and N₂O exchanges between soils and the atmosphere play an important role in climate warming and global carbon and nitrogen cycling; however, it is still not clear whether the fluxes of these two greenhouse gases are correlated at the ecosystem scale. We collected 143 pairs of ecosystem CO₂ and N₂O exchanges between soils and the atmosphere measured simultaneously in eight ecosystems around the world and developed relationships between soil CO₂ and N₂O fluxes. Significant linear regressions of soil CO₂ and N₂O fluxes were found for all eight ecosystems; the highest slope occurred in rice paddies and the lowest in temperate grasslands. We also found the dominant role of growing season on the relationship of annual CO₂ and N₂O fluxes. No significant relationship between soil CO₂ and N₂O fluxes was found across all eight ecosystem types. The estimated annual global N₂O emission based on our findings is 13.31 Tg N yr⁻¹ with a range of 8.19–18.43 Tg N yr⁻¹ for 1980–2000, of which cropland contributes nearly 30%. Our findings demonstrated that stoichiometric relationships may work on ecological functions at the ecosystem level. The relationship of soil N₂O and CO₂ fluxes developed here could be helpful in biogeochemical modeling and large-scale estimations of soil CO₂ and N₂O fluxes.     

Carbon stored in human settlements: the conterminous United States

Urban areas are home to more than half of the world's people, responsible for >70% of anthropogenic release of carbon dioxide and 76% of wood used for industrial purposes. By 2050 the proportion of the urban population is expected to increase to 70% worldwide. Despite fast rates of change and potential value for mitigation of carbon dioxide emissions, the organic carbon storage in human settlements has not been well quantified. Here, we show that human settlements can store as much carbon per unit area (23–42 kg C m⁻² urban areas and 7–16 kg C m⁻²exurban areas) as tropical forests, which have the highest carbon density of natural ecosystems (4–25 kg C m⁻²). By the year 2000 carbon storage attributed to human settlements of the conterminous United States was 18 Pg of carbon or 10% of its total land carbon storage. Sixty-four percent of this carbon was attributed to soil, 20% to vegetation, 11% to landfills, and 5% to buildings. To offset rising urban emissions of carbon, regional and national governments should consider how to protect or even to increase carbon storage of human-dominated landscapes. Rigorous studies addressing carbon budgets of human settlements and vulnerability of their carbon storage are needed.     

On carbon sequestration in desert ecosystems

Recent reports of net ecosysytem production >100 gCm⁻² yr⁻¹ in deserts are incompatible with existing measurements of net primary production and carbon pools in deserts. The comparisions suggest that gas exchange measurements should be used with caution and better validation if they are expected to indicate the magnitude of carbon sink in these ecosysytems.     

Nitrogen effects on net ecosystem carbon exchange in a temperate steppe

It has widely been documented that nitrogen (N) enrichment stimulates plant growth and net primary production. However, there is still dispute on how N addition affects net ecosystem CO₂ exchange (NEE), which represents the balance between ecosystem carbon (C) uptake and release. We conducted an experimental study to examine effects of N addition on NEE in a temperate steppe in northern China from 2005 to 2008. N was added at a rate of 10 g N m⁻² yr⁻¹ with NH₄NO₃ alone or in combination with phosphorous (P, 5 g P₂O₅ m⁻² yr⁻¹) in both clipped and unclipped plots. Over the 4 years, N addition significantly stimulated growing-season NEE, on average, by 27%. Neither the main effects of P addition or clipping nor their interactions with N addition were statistically significant on NEE in any of the 4 years. However, the magnitude of N stimulation on NEE declined over time. N addition significantly increased NEE by 60% in 2005 and 21% in 2006, but its effect was not significant in 2007 and 2008. N-induced shift in species composition was primarily responsible for the declined N stimulation over time. The gradually increasing coverage of the upper canopy species ( Stipa krylovii ) and standing litter accumulation induced light limitation on the lower canopy species ( Artemisia frigida ). Thus, N-induced shifts in plant species composition strongly regulated the direct effects of N addition on C sequestration in the temperate steppe.     

Population dynamics and production of the pelagic amphipod Hyalella montezuma in a thermally constant system

SUMMARY. 1. The population dynamics and annual production of the multivoltine. pelagic amphipod Hyalella montezuma were studied over a 3-year period in the thermally constant environment of Montezuma Well. Arizona.
2. H. montezuma showed two maxima which coincided with spring and autumn phytoplankton maxima. Juveniles comprised over 85% of the population in the pelagic zone compared to 37% in the littoral vegetation and there were significantly more females in the littoral vegetation. It appears that juvenile and adult H. montezuma show different habitat preferences.
3. Total annual mean production for H. montezuma calculated by the size frequency method and adjusted for multiple generations, was 357 kg ha⁻¹ yr⁻¹, which is higher than single-species production estimates reported for most zoobenthic amphipods and multivoltine planktonic crustaceans. Average energy production for H. montezuma was 4640 kJ ha⁻¹ yr⁻¹ in the pelagic zone and 1072 kJ ha⁻¹ yr⁻¹ in the littoral vegetation.
4. Average cohort P / ratios for H. montezuma were higher in the pelagic zone (5.5) than in the litttoral vegetation (3.7). Juveniles had higher cohort P ratios than adults in the pelagic zone, while the reverse relationship was true in the littoral vegetation. We propose that different size-selective predators may contribute to the differences in P ratios for juveniles and adults in these two habitats.