Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

Long-term effects of nitrogen fertilization on the productivity of subsequent stands of Douglas-fir in the Pacific Northwest

The carryover effects of N fertilization on five coastal Pacific Northwest Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) plantations were studied. “Carryover” is defined as the long-term impact of N fertilizer added to a previous stand on the growth of a subsequent stand. Average height and diameter at 1.3 m above-ground (DBH) of 7–9-year-old Douglas-fir trees and biomass and N-content of understory vegetation were assessed on paired control (untreated) and urea-N-fertilized plots that had received cumulative additions of 810–1120 kg N ha⁻¹ to a previous stand. Overall productivity was significantly greater in the fertilized stands compared to the controls. In 2006, the last growth measurement year, mean seedling height was 15% greater (p = 0.06) and mean DBH was 29% greater (p = 0.04) on previously fertilized plots compared to control plots. Understory vegetation biomass of fertilized plots was 73% greater (p = 0.005), and N-content was 97% greater (p = 0.004) compared to control plots. These results show that past N fertilization markedly increased seedling growth in these plantations as well as biomass and N-content of understory vegetation in a subsequent rotation. These findings suggest that N fertilization could potentially increase site productivity of young Douglas-fir stands found on low quality sites in the Pacific Northwest 15–22 years after application by a carryover effect. These plantations have not yet reached the age where marketable materials can be harvested from them, and the growth of trees should be monitored over a longer time period before potential impacts on older stands, if any, can be determined.     

Recovery of carbon and nutrient pools in a northern forested wetland 11 years after harvesting and site preparation

We measured the change in above- and below-ground carbon and nutrient pools 11 years after the harvesting and site preparation of a histic-mineral soil wetland forest in the Upper Peninsula of Michigan. The original stand of black spruce (Picea mariana), jack pine (Pinus banksiana) and tamarack (Larix laricina) was whole-tree harvested, and three post-harvest treatments (disk trenching, bedding, and none) were randomly assigned to three Latin square blocks (n = 9). Nine control plots were also established in an adjoining uncut stand. Carbon and nutrients were measured in three strata of above-ground vegetation, woody debris, roots, forest floor, and mineral soil to a depth of 1.5 m. Eleven years following harvesting, soil C, N, Ca, Mg, and K pools were similar among the three site preparation treatments and the uncut stand. However, there were differences in ecosystem-level nutrient pools because of differences in live biomass. Coarse roots comprised approximately 30% of the tree biomass C in the regenerated stands and 18% in the uncut stand. Nutrient sequestration, in the vegetation since harvesting yielded an average net ecosystem gain of 332 kg N ha⁻¹, 110 kg Ca ha⁻¹, 18 kg Mg ha⁻¹, and 65 kg K ha⁻¹. The likely source for the cations and N is uptake from shallow groundwater, but N additions could also come from non-symbiotic N-fixation and N deposition. These are the only reported findings on long-term effects of harvesting and site preparation on a histic-mineral soil wetland and the results illustrate the importance of understanding the ecohydrology and nutrient dynamics of the wetland forest. This wetland type appears less sensitive to disturbance than upland sites, and is capable of sustained productivity under these silvicultural treatments.     

Growth and nutrition of Pinus radiata in response to fertilizer applied after thinning and interaction with defoliation associated with Essigella californica

Infestations of Essigella californica following the installation of post-thinning fertilizer trials in Pinus radiata plantations provided an opportunity to examine the impact of repeated defoliation over a period of 8 years (1997–2005). Replicated treatments (n = 4) of nil fertilizer (control), N (300 kg ha⁻¹) as urea, P (80 kg ha⁻¹) and S (45 kg ha⁻¹) as superphosphates were applied immediately after thinning at three sites and this was followed by a second application of NPS fertilizers 6 years later with N applied at 300 kg ha⁻¹ as urea and ammonium sulphate and P at 80 or 120 kg ha⁻¹. Defoliation of untreated P. radiata gradually increased to 50% over a period of 8 years. Basal area growth was negatively correlated with average defoliation for two consecutive post-fertilizer periods of 6 and 2 years. Growth responses to fertilizer varied considerably between sites but the largest improvement in growth was due to NPS fertilizer, this increased basal area by 30–80%. Application of N fertilizer raised total N levels in foliage and increased defoliation with a commensurate loss in growth under conditions of deficiencies of S or P. Repeated infestations gradually increased the percentage of trees with severe defoliation (>80% loss of foliage) indicating that nutrient-deficient trees have a reduced capacity for foliage recovery between episodes of peak infestation. In contrast, treatment with N fertilizer in combination with S- and P-corrected deficiencies of these nutrients, raised levels of total N in foliage and reduced defoliation to approximately 20%. Basal area growth responses to NPS fertilizers reflected improved nutrition as well as reduced insect damage. The reduction in defoliation under conditions of balanced tree nutrition was most likely due to enhanced needle retention following correction of P deficiency as well as greater availability of nutrients enabling a more vigorous recovery of P. radiata after an episode of E. californica activity. Treatment with fertilizer therefore reduced the long-term impact of aphid damage and improved growth of P. radiata.     

Key nitrogen cycling processes in pine plantations along a short urban–rural gradient in Nanchang,China

We used pine (Pinus elliottii Engelm.) forests located along a short urban–rural gradient in Nanchang, China to study nitrogen (N) cycling responses to urbanization. Annual average rates of nitrification and net N-mineralization in soils (0–15 cm depth) measured from February 2007 to January 2009 increased from rural (8 and 37 kg ha⁻¹ year⁻¹) to suburban (69 and 79 kg ha⁻¹ year⁻¹) and urban sites (114 and 116 kg ha⁻¹ year⁻¹) (P < 0.05). Soil nitrate and mineral N pools exhibited the same spatial patterns in response to urban location. In comparison to rural sites, urban and suburban sites experienced soil microbial biomass N that increased by 98% and 38%, sucrase activity that increased by 40% and 26%, and urease activity that decreased by 35% and 25%, respectively. Soil microbial biomass C:N and free amino acids varied little along the urban–rural gradient. Foliar N concentrations and N resorption proficiencies were higher in urban (12.3 and 4.8 g kg⁻¹) and suburban (12.3 and 6.2 g kg⁻¹) than in rural (9.9 and 3.6 g kg⁻¹) sites, while N resorption efficiencies (from 58% to 72%) were not statistically different. These results indicate that forests in suburban and especially in urban areas are moving rapidly towards a state of “N saturation” and increased potential N loss most likely attributable to higher N deposition to these sites.     

Above- and belowground biomass in a Brazilian Cerrado

Cerrado is a biome that occupies about 25% of the Brazilian territory and is characterized by a gradient of grassland to savanna and forest formations and by high species richness. It has been severely affected by degradation and deforestation and has been heavily fragmented over the past 4-5 decades. Despite the recognized overall ecological importance of the Cerrado, there are only few studies focusing on the quantification of biomass in this biome. We conducted such a case study in the South-East of Brazil in a cerrado sensu stricto (cerrado s.s.) with the goal to produce estimates of above- and belowground biomass and to develop allometric equations. A number of 120 trees from 18 species were destructively sampled and partitioned into the components: leaves, branches and bole. Five models with DBH (D), height (H), D²H and wood density (WD) as independent variables were tested for the development of allometric models for individual tree aboveground biomass (leaves + branches + bole). One model based on basal area (BA) as a stand parameter was also tested as an alternative approach for predicting aboveground biomass in the stand level. Belowground biomass was estimated by subsampling on 10 sample plots. Mean aboveground tree biomass (bole, branches and leaves) was estimated to be 62,965.5 kg ha⁻¹(SE = 14.6%) and belowground biomass accounted for 37,501.8 kg ha⁻¹ (SE = 23%). The best-fit equation for the estimation of individual tree aboveground biomass include DBH and wood density as explanatory variables (R² = 0.898; SEE = 0.371) and is applicable for the diameter range of this study (5.0-27.6 cm) and in environments with similar conditions of the cerrado s.s. sampled. In the stand level, the model tested presented a higher goodness of fit than the single tree models (R² = 0.934; SEE = 0.224). Our estimates of aboveground biomass are higher than reported by other studies developed in the same physiognomy, but the estimates of belowground biomass are within the range of values reported in other studies from sites in cerrado s.s. Both biomass estimates, however, exhibit relatively large standard errors. The root-to-shoot ratio of the sample trees is in the magnitude of reported values for savanna ecosystems, but smaller than estimated from other studies in the cerrado s.s.     

Specific gravity responses of slash and loblolly pine following mid-rotation fertilization

Wood quality attributes were examined in six stands of slash pine (Pinus elliottii Engelm. var. elliottii) and loblolly pine (P. taeda L.) in the lower Coastal Plain of Georgia and Florida. Several plots comprised each stand, and each plot was divided so that it received three fertilizer treatments: a control treatment with herbaceous weed control at planting and brush control at mid-rotation only (control); 45 kg ha⁻¹ N + 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer with N at planting); and 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer without N at planting). Ring width, ring earlywood specific gravity (SG), ring latewood SG, whole ring SG, and ring percent latewood were measured on each of seven trees. Of these measurements, this study focused mainly on the properties related to SG. Examination of the rings showed that latewood SG was significantly lower in trees treated with fertilizers with and without N at planting in the two to three years following fertilization, but that latewood SG gradually returned to a level similar to the control. Fertilizer without N at planting may also have had a brief negative effect on earlywood SG following mid-rotation fertilization, but it was not as clear or lasting as the effect on latewood SG. Additionally, although slash and loblolly pine appear to differ in the developmental patterns of these SG properties, there were no significant differences in how these patterns interacted with treatment. This study demonstrated that fertilization treatments have similar short-term effects on the SG of slash and loblolly pines, particularly in latewood, but the trees will return to a SG pattern consistent with unfertilized trees within two or three years.     

Carbon storage in old-growth and second growth fire-dependent western larch (Larix occidentalis Nutt.) forests of the Inland Northwest,USA

There is limited understanding of the carbon (C) storage capacity and overall ecological structure of old-growth forests of western Montana, leaving little ability to evaluate the role of old-growth forests in regional C cycles and ecosystem level C storage capacity. To investigate the difference in C storage between equivalent stands of contrasting age classes and management histories, we surveyed paired old-growth and second growth western larch (Larix occidentalis Nutt)–Douglas-fir (Pseudostuga menziesii var. glauca) stands in northwestern Montana. The specific objectives of this study were to: (1) estimate ecosystem C of old-growth and second growth western larch stands; (2) compare C storage of paired old-growth–second growth stands; and (3) assess differences in ecosystem function and structure between the two age classes, specifically measuring C associated with mineral soil, forest floor, coarse woody debris (CWD), understory, and overstory, as well as overall structure of vegetation. Stands were surveyed using a modified USFS FIA protocol, focusing on ecological components related to soil, forest floor, and overstory C. All downed wood, forest floor, and soil samples were then analyzed for total C and total nitrogen (N). Total ecosystem C in the old-growth forests was significantly greater than that in second growth forests, storing over 3 times the C. Average total mineral soil C was not significantly different in second growth stands compared to old-growth stands; however, total C of the forest floor was significantly greater in old-growth (23.8 Mg ha⁻¹) compared to second growth stands (4.9 Mg ha⁻¹). Overstory and coarse root biomass held the greatest differences in ecosystem C between the two stand types (old-growth, second growth), with nearly 7 times more C in old-growth trees than trees found on second growth stands (144.2 Mg ha⁻¹ vs. 23.8 Mg ha⁻¹). Total CWD on old-growth stands accounted for almost 19 times more C than CWD found in second growth stands. Soil bulk density was also significantly higher on second growth stands some 30+ years after harvest, demonstrating long-term impacts of harvest on soil. Results suggest ecological components specific to old-growth western larch forests, such as coarse root biomass, large amounts of CWD, and a thick forest floor layer are important contributors to long-term C storage within these ecosystems. This, combined with functional implications of contrasts in C distribution and dynamics, suggest that old-growth western larch/Douglas-fir forests are both functionally and structurally distinctive from their second growth counterparts.     

Above- and belowground nutrients storage and biomass accumulation in marginal Nothofagus antarctica forests in Southern Patagonia

The above- and belowground biomass and nutrient content (N, P, K, Ca, S and Mg) of pure deciduous Nothofagus antarctica (Forster f.) Oersted stands grown in a marginal site and aged from 8 to 180 years were measured in Southern Patagonia. The total biomass accumulated ranged from 60.8 to 70.8 Mg ha⁻¹ for regeneration and final growth stand, respectively. The proportions of belowground components were 51.6, 47.2, 43.9 and 46.7% for regeneration, initial growth, final growth and mature stand, respectively. Also, crown classes affected the biomass accumulation where dominant trees had 38.4 Mg ha⁻¹ and suppressed trees 2.6 Mg ha⁻¹ to the stand biomass in mature stand. Nutrient concentrations varied according to tree component, crown class and stand age. Total nutrient concentration graded in the fallowing order: leaves > bark > middle roots > small branches > fine roots > sapwood > coarse roots > heartwood. While N and K concentrations increased with age in leaves and fine roots, concentration of Ca increased with stand age in all components. Dominant trees had higher N, K and Ca concentrations in leaves, and higher P, K and S concentrations in roots, compared with suppressed trees. Although the stands had similar biomass at different ages, there were important differences in nutrient accumulation per hectare from 979.8 kg ha⁻¹ at the initial growth phase to 665.5 kg ha⁻¹ at mature stands. Nutrient storage for mature and final growth stands was in the order Ca > N > K > P > Mg > S, and for regeneration stand was Ca > N > K > Mg > P > S. Belowground biomass represented an important budget of all nutrients. At early ages, N, K, S, Ca and Mg were about 50% in the belowground components. However, P was 60% in belowground biomass and then increased to 70% in mature stands. These data can assist to quantify the impact of different silviculture practices which should aim to leave material (mainly leaves, small branches and bark) on the site to ameliorate nutrient removal and to avoid a decline of long-term yields.     

Carbon density and accumulation in woody species of tropical dry forest in India

We studied the carbon density and accumulation in trees at five sites in a tropical dry forest (TDF) to address the questions: how is the TDF structured in terms of tree and carbon density in different DBH (diameter at breast height) classes? What are the levels of carbon density and accumulation in the woody species of TDF? Is the vegetation carbon density evenly distributed across the forest? Does carbon stored in the soil reflect the pattern of aboveground vegetation carbon density? Which species in the forest have a high potential for carbon accumulation? The WSG among species ranged from 0.39 to 0.78 g cm⁻³. Our study indicated that most of the carbon resides in the old-growth (high DBH) trees; 88-97% carbon occurred in individuals ?19.1 cm DBH, and therefore extra care is required to protect such trees in the dry forest. Acacia catechu, Buchanania lanzan, Hardwickia binata, Shorea robusta and Terminalia tomentosa accounted for more than 10 t ha⁻¹ carbon density, warranting extra efforts for their protection. Species also differed in their capacity to accumulate carbon indicating variable suitability for afforestation. Annually, the forest accumulated 5.3 t-C ha⁻¹ yr⁻¹ on the most productive, wettest Hathinala site to 0.05 t-C ha⁻¹ yr⁻¹ on the least productive, driest Kotwa site. This study indicated a marked patchy distribution of carbon density (151 t-C ha⁻¹ on the Hathinala site to 15.6 t-C ha⁻¹ on the Kotwa site); the maximum value was more than nine times the minimum value. These findings suggest that there is a substantial scope to increase the carbon density and accumulation in this forest through management strategies focused on the protection, from deforestation and fire, of the high carbon density sites and the old-growth trees, and increasing the stocking density of the forest by planting species with high potential for carbon accumulation.     

Soil changes induced by Acacia mangium plantation establishment: Comparison with secondary forest and Imperata cylindrica grassland soils in South Sumatra,Indonesia

Acacia plantation establishment might cause soil acidification in strongly weathered soils in the wet tropics because the base cations in the soil are translocated rapidly to plant biomass during Acacia growth. We examined whether soils under an Acacia plantation were acidified, as well as the factors causing soil acidification. We compared soils from 10 stands of 8-year-old Acacia mangium plantations with soils from 10 secondary forests and eight Imperata cylindrica grasslands, which were transformed into Acacia plantations. Soil samples were collected every 5–30 cm in depth, and pH and related soil properties were analyzed. Soil pH was significantly lower in Acacia plantations and secondary forests than in Imperata grasslands at every soil depth. The difference was about 1.0 pH unit at 0–5 cm and 0.5 pH unit at 25–30 cm. A significant positive correlation between pH and base saturation at 0–20 cm depth indicated that the low pH under forest vegetation was associated with exchangeable cation status. Using analysis of covariance (ANCOVA), with clay content as the covariate, exchangeable Ca (Ex-Ca) and Mg (Ex-Mg) stocks were significantly lower in forested areas than in Imperata grasslands at any clay content which was strongly related to exchangeable cation stock. The adjusted average Ex-Ca stock calculated by ANCOVA was 249 kg ha⁻¹ in Acacia plantations, 200 kg ha⁻¹ in secondary forests, and 756 kg ha⁻¹ in Imperata grasslands at 0–30 cm. Based on a comparison of estimated nutrient stocks in biomass and soil among the vegetation types, the translocation of base cations from soil to plant biomass might cause a decrease in exchangeable cations and soil acidification in Acacia plantations.     

Formulating allometric equations for estimating biomass and carbon stock in small diameter trees

Biomass and carbon sequestration rate of a young (four year old) mixed plantation of Dalbergia sissoo Roxb., Acacia catechu Willd., and Albizia lebbeck Benth. growing in Terai region (a level area of superabundant water) of central Himalaya was estimated. The plantation is seed sown in the rainy season of year 2004 and spread over an area of 44 ha. Allometric equations for both above and below ground components were developed for three tree species. The density of trees in the plantation was 1322 trees ha⁻¹ The diameters of trees were below 10 cm. Five diameter classes were defined for D. sissoo and A. catechu and 3 for A. lebbeck. 5 trees were harvested in each diameter class. Individual tree allometry was exercised for developing the allometric equations relating tree component (low and above ground) biomass to d.b.h. Post analysis equations were highly significant (P > 0.001) for each component of all species. In the plantation Holoptelia integrifolia Roxb. (Family Ulmaceae) has been reduced to shrub form because of frost. Only the aboveground biomass of H. integrifolia and other shrubs were estimated by destructive harvesting method. Herbaceous forest floor biomass and leaf litter fall were also estimated. The total forest vegetation biomass was 10.86 Mg ha⁻¹ in 2008 which increased to 19.49 Mg ha⁻¹ in 2009. The forest is sequestering carbon at the rate of 4.32 Mg ha⁻¹ yr⁻¹.     

Carbon accumulation in the biomass and soil of different aged secondary forests in the humid tropics of Costa Rica

Efforts are needed in order to increase confidence for carbon accounts in the land use sector, especially in tropical forest ecosystems that often need to turn to default values given the lack of precise and reliable site specific data to quantify their carbon sequestration and storage capacity. The aim of this study was then to estimate biomass and carbon accumulation in young secondary forests, from 4 and up to 20 years of age, as well as its distribution among the different pools (tree including roots, herbaceous understory, dead wood, litter and soil), in humid tropical forests of Costa Rica. Carbon fraction for the different pools and tree components (stem, branches, leaves and roots) was estimated and varies between 37.3% (±3.3) and 50.3% (±2.9). Average carbon content in the soil was 4.1% (±2.1). Average forest plant biomass was 82.2 (±47.9) Mg ha⁻¹ and the mean annual increment for carbon in the biomass was 4.2 Mg ha⁻¹ yr⁻¹. Approximately 65.2% of total biomass was found in the aboveground tree components, while 14.2% was found in structural roots and the rest in the herbaceous vegetation and necromass. Carbon in the soil increased by 1.1 Mg ha⁻¹ yr⁻¹. Total stored carbon in the forest was 180.4 Mg ha⁻¹ at the age of 20 years. In these forests, most of the carbon (51-83%) was stored in the soil. Models selected to estimate biomass and carbon in trees as predicted by basal area had R² adjustments above 95%. Results from this study were then compared with those obtained for a variety of secondary and primary forests in different Latin-American tropical ecosystems and in tree plantations in the same study area.     

Comparison of 5th- and 14th-year Douglas-fir and understory vegetation responses to selective vegetation removal

The effects of early vegetation management on the survival and growth of Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] were examined 5 and 15 years after planting in the Oregon Coast Range. Our first objective was to document the effects of vegetation species competition upon key ecosystem properties. The second objective was to document the effects of vegetation removal during early Douglas-fir stand establishment upon long-term tree growth and on biomass production by vegetation components. Seven levels of manual vegetation removal were maintained for the first 5 years after planting: 0%, 25%, 50%, 75%, and 100% shrub removal; and 100% shrub removal combined with 50% or 100% herbaceous vegetation removal. Shrub and herb removal did not affect Douglas-fir survival at year five, but treatments providing less than 75% shrub removal significantly reduced Douglas-fir survival by year 15. Removing shrubs and herbs completely (100S + 100H) during the 5 years following tree planting allowed successful tree establishment, with a 366% increase in biomass accumulation per hectare for Douglas-fir in that treatment at the end of 14 years of growth. At 15 years stand age, even with shrub removal alone, a 304% gain in tree biomass per hectare was obtained compared to no vegetation removal (NVR). By stand age 15 years, any increase in the degree of understory removal beyond 75% did not contribute significantly to additional tree survival and growth. The understory vegetation on NVR treatment plots and the herbaceous vegetation on 100% shrub removal (100S) treatment plots, contained >90% and >80% of aboveground biomass N at 5 years, respectively, indicating possible competition for soil N. Soil moisture was not different among treatments at 5 years. Complete vegetation removal (100S + 100H) for 5 years resulted in a significant increase in soil bulk density (P < 0.05), a significant decrease in total soil C (P < 0.05) and no change in total soil N in the upper 15 cm of the mineral soil. By 14 years, however, only the soil bulk density remained greater (P < 0.05) on the 100S + 100H treatment. We conclude that greater tree survival and growth occurred with at least 75% shrub removal. Our results suggest that managers may have substantial flexibility in maintaining a partial understory component suitable for ecosystem productivity, canopy cover and wildlife habitat, while maintaining forests productive for timber resources.     

A comparison of annual transpiration and productivity in monoculture and mixed-species Douglas-fir and red alder stands

Although much is known about drivers of productivity in Douglas-fir and red alder stands, less is known about how productivity may relate to stand transpiration and water use efficiency. We took advantage of a 15-year-old experiment involving Douglas-fir (Pseudotsuga menziesii) and red alder (Alnus rubra) in the western Cascade Range of western Oregon to test the following hypotheses: (a) more productive stands transpire more water, (b) the relationship between productivity and transpiration differs between species, and (c) the relationship between productivity and transpiration differs between sites varying in soil moisture and fertility. Furthermore, the experimental design included alder, a facultative nitrogen-fixing species, which could also affect fertility. Fixed area plots (20 × 20 m) were planted as monocultures of each species or in mixtures at a common density (1100 trees ha⁻¹) in a randomized-block design. Transpiration of Douglas-fir and red alder was measured using heat dissipation sensors installed in eight trees per plot and scaled to the plot level based on sapwood basal area for each species. Although up to 53% of the variability in tree transpiration was explained by basal area, irrespective of species or site conditions, the two stands with the highest biomass and sapwood basal area did not transpire the most. Instead of more productive stands transpiring more water, the greatest variability in both productivity and transpiration was determined by site conditions and to a lesser degree, species composition. For example, 70% of the variation in tree biomass increment (TBI) was determined by leaf area index, which was much higher at the site with higher fertility and soil moisture (p < 0.05). Despite marked phenological and physiological differences, Douglas-fir and red alder performed similarly. Only 19% of annual water use of Douglas-fir occurred between October and March when alder was leafless. Also, there was no evidence of a fertilization effect of the nitrogen-fixing red alder on the Douglas-fir: the nitrogen concentration and N-isotopic ratio of Douglas-fir needles did not differ whether trees were grown in monoculture or in mixtures with red alder. We conclude that lower soil fertility and contrasting microclimate at one site relative to the other suppressed NPP while maintaining higher transpiration, thus reducing water use efficiency.     

Ecosystem production in Douglas-fir plantations: Interaction of red alder and site fertility

Pure stands of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and mixed stands of Douglas-fir and naturally established red alder (Alnus rubra Bong.) were examined on two different sites for soil properties, tree growth and nutrition, and aboveground ecosystem biomass and net primary production. One site (Mt. Benson, Nanaimo, B.C.) was nitrogen (N)-deficient and had a low site index (expected Douglas-fir height of 24 m at 50 years). The other site (Skykomish, western Washington) was N-rich and had a site index of 45 m at 50 years. Soil N accretion on the red alder units was estimated at 65 (Mt. Benson) and 42 (Skykomish) kg ha^?1 year^?1 for 23 years to a soil depth of 50 cm. At the current stage of plantation development, presence of red alder at the infertile Mt. Benson site increased average Douglas-fir diameter but did not affect its basal area and basal area growth rate; including alder stem biomass increased total stand basal area and basal area growth 2.5 fold. Presence of red alder at the fertile Skykomish site decreased average diameter, basal area, and basal area growth of Douglas-fir; including alder biomass left total stand basal area and basal area growth unchanged. Douglas-fir foliar N concentrations on Mt. Benson increased from 0.93 without alder to 1.41% on the red alder unit but were 1.55% for both units at Skykomish. Although alder did not affect Douglas-fir aboveground biomass and net primary production on Mt. Benson, total ecosystem biomass doubled and production tripled when alder biomass was included. Conversely, at Skykomish, Douglas-fir biomass and production decreased, and total ecosystem values were essentially unchanged. Mixing red alder and Douglas-fir seems to have great potential for increasing Douglas-fir growth and ecosystem production on infertile, N-deficient sites but probably has limited value on fertile, N-rich sites.     

An ecosystem approach to biodiversity effects: Carbon pools in a tropical tree plantation

This paper presents a synthesis of experiments conducted in a tropical tree plantation established in 2001 and consisting of 22 plots of 45 m × 45 m with either one, three or six native tree species. We examined the changes in carbon (C) pools (trees, herbaceous vegetation, litter, coarse woody debris (CWD), and mineral topsoil at 0-10 cm depth) and fluxes (decomposition of CWD and litter, as well as soil respiration) both through time and among diversity levels. Between 2001 and 2009 the aboveground C pools increased, driven by trees. Across diversity levels, the mean observed aboveground C pool was 7.9 ± 2.5 Mg ha⁻¹ in 2006 and 20.4 ± 7.4 Mg ha⁻¹ in 2009, a 158% increase. There was no significant diversity effect on the observed aboveground C pool, but we found a significant decrease in the topsoil C pool, with a mean value of 34.5 ± 2.4 Mg ha⁻¹ in 2001 and of 25.7 ± 5.7 Mg ha⁻¹ in 2009 (F_1,36 = 52.12, p < 0.001). Assuming that the biomass C pool in 2001 was negligible (<1 Mg ha⁻¹), then the plantation gained in C, on average, ∼20 and lost ∼9 Mg ha⁻¹ in biomass and soil respectively, for an overall gain of ∼11 Mg ha⁻¹ over 8 years. Across the entire data set, we uncovered significant effects of diversity on CWD decomposition (diversity: F_2,393 = 15.93, p < 0.001) and soil respiration (monocultures vs mixtures: t = 15.35, df = 11, p < 0.05) and a marginally significant time × diversity interaction on the loss of total C from the mineral topsoil pool (see above). Monthly CWD decomposition was significantly faster in monocultures (35.0 ± 24.1%) compared with triplets (31.3 ± 21.0%) and six-species mixtures (31.9 ± 26.8%), while soil respiration was higher in monocultures than in mixtures (t = 15.35, df = 11, p < 0.001). Path analyses showed that, as diversity increases, the links among the C pools and fluxes strengthen significantly. Our results demonstrate that tree diversity influences the processes governing the changes in C pools and fluxes following establishment of a tree plantation on a former pasture. We conclude that the choice of tree mixtures for afforestation in the tropics can have a marked influence on C pools and dynamics.     

Nitrogen leaching in response to increased nitrogen inputs in subtropical monsoon forests in southern China

Dissolved inorganic nitrogen (DIN) (as ammonium nitrate) was applied monthly onto the forest floor of one old-growth forest (>400 years old, at levels of 50, 100 and 150 kg N ha⁻¹ yr⁻¹) and two young forests (both about 70 years old, at levels of 50 and 100 kg N ha⁻¹ yr⁻¹) over 3 years (2004–2006), to investigate how nitrogen (N) input influenced N leaching output, and if there were differences in N retention between the old-growth and the young forests in the subtropical monsoon region of southern China. The ambient throughfall inputs were 23–27 kg N ha⁻¹ yr⁻¹ in the young forests and 29–35 kg N ha⁻¹ yr⁻¹ in the old-growth forest. In the control plots without experimental N addition, a net N retention was observed in the young forests (on average 6–11 kg N ha⁻¹ yr⁻¹), but a net N loss occurred in the old-growth forest (−13 kg N ha⁻¹ yr⁻¹). Experimental N addition immediately increased DIN leaching in all three forests, with 25–66% of added N leached over the 3-year experiment. At the lowest level of N addition (50 kg N ha⁻¹ yr⁻¹), the percentage N loss was higher in the old-growth forest (66% of added N) than in the two young forests (38% and 26%). However, at higher levels of N addition (100 and 150 kg N ha⁻¹ yr⁻¹), the old-growth forest exhibited similar N losses (25–43%) to those in the young forests (28–43%). These results indicate that N retention is largely determined by the forest successional stages and the levels of N addition. Compared to most temperate forests studied in Europe and North America, N leaching loss in these seasonal monsoon subtropical forests occurred mainly in the rainy growing season, with measured N loss in leaching substantially higher under both ambient deposition and experimental N additions.     

Estimating biomass production and carbon storage for a fast-growing makino bamboo (Phyllostachys makinoi) plant based on the diameter distribution model

The purpose of this study was to estimate biomass and carbon storage for a fast-growing makino bamboo (Phyllostachys makinoi). The study site was located in central Taiwan and the makino bamboo plantation had a stand density of 21191 ± 4107 culms ha⁻¹. A diameter distribution model based on the Weibull distribution function and an allometric model was used to predict aboveground biomass and carbon storage. For an accurate estimation of carbon storage, the percent carbon content (PCC) in different sections of bamboo was determined by an elemental analyzer. The results showed that bamboos of all ages shared a similar trend, where culms displayed a carbon storage of 47.49–47.82%, branches 45.66–46.23%, and foliage 38.12–44.78%. In spite of the high density of the stand, the diameter distribution of makino bamboo approached a normal distribution and aboveground biomass and carbon storage were 105.33 and 49.81 Mg ha⁻¹, respectively. Moreover, one-fifth of older culms from the entire stand were removed by selective cutting. If the distribution of the yield of older culms per year was similar to the current stand, the yields of biomass and carbon per year would be 21.07 and 9.89 Mg ha⁻¹ year⁻¹. An astonishing productivity was observed, where every 5 years the yield of biomass and carbon was equal to the current status of stockings. Thus, makino bamboo has a high potential as a species used for carbon storage.