Growth, foliar nutrition and δC responses of red alder (Alnus rubra) to phosphorus additions soon after planting on moist sites

In south-coastal British Columbia, a low availability of phosphorus (P) may limit the early growth of young red alder (Alnus rubra Bong.), even on sites classified as productive for red alder. However, it remains unclear as to what P addition rates best alleviate P deficiencies on such sites and how long effects of P additions on growth persist.We applied P 1-3 months after planting at rates up to 60 g P tree⁻¹ and assessed growth and foliar elemental contents over three growing seasons at three sites with site productivity classed as good for red alder. Foliar δ¹³C was also determined in year 1 in the two sites on Vancouver Island and in year 2 in the site on the British Columbia mainland coast in order to better understand the relationships among foliar nutritional status, leaf water use efficiency (WUE), and growth.P additions at planting significantly increased height (11-15%), diameter (26%) and stem volume (62-64%) through 3 years. Maximum growth rates were achieved at P addition rates of 30 g tree⁻¹ and at foliar P concentrations of 2.2-2.5 g kg⁻¹. Growth did not increase further at addition rates of 60 g P tree⁻¹. Stem growth increases were accompanied by increased individual leaf mass, first-year foliar concentrations of N, P, Ca, Mg, and S, and foliar δ¹³C, the latter suggesting that WUE increased with P additions. Foliar concentrations of P in unfertilized trees were at deficient levels, based on earlier studies, and increases in first-year foliar P concentrations and stem growth through year 3 were consistent with responses in earlier single-tree plot experiments. Longer-term measurements are required to define the duration of growth response to P additions in these otherwise-productive sites.     

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.     

Effects of [CO2] and nitrogen on morphological and biomass traits of white birch (Betula papyrifera) seedlings

To investigate the interactive effects of CO₂ concentration ([CO₂]) and nitrogen supply on the growth and biomass of boreal trees, white birch seedlings (Betula papyrifera) were grown under ambient (360 μmol mol⁻¹) and elevated [CO₂] (720 μmol mol⁻¹) with five nitrogen supply regimes (10, 80, 150, 220, and 290 μmol mol⁻¹) in greenhouses. After 90 days of treatment, seedling height, root-collar diameter, biomass of different organs, leaf N concentration, and specific leaf area (SLA) were measured. Significant interactive effects of [CO₂] and N supply were found on height, root-collar diameter, leaf biomass, stem biomass and total biomass, stem mass ratio (SMR), and root mass ratio (RMR), but not on root mass, leaf mass ratio (LMR), leaf to root ratio (LRR), or leaf N concentration. The CO₂ elevation generally increased all the growth and biomass parameters and the increases were generally greater at higher levels of N supply or higher leaf N concentration. However, the CO₂ elevation significantly reduced SLA (13.4%) and mass-based leaf N concentration but did not affect area-based leaf N concentration. Increases in N supply generally increased the growth and biomass parameters, but the relationships were generally curvilinear. Based on a second order polynomial model, the optimal leaf N concentration was 1.33 g m⁻² for height growth under ambient [CO₂] and 1.52 g m⁻² under doubled [CO₂]; 1.48 g m⁻² for diameter under ambient [CO₂] and 1.64 g m⁻² under doubled [CO₂]; 1.29 g m⁻² for stem biomass under ambient [CO₂] and 1.43 g m⁻² under doubled [CO₂]. The general trend is that the optimal leaf N was higher at doubled than ambient [CO₂]. However, [CO₂] did not affect the optimal leaf N for leaf and total biomass. The CO₂ elevation significantly increased RMR and SMR but decreased LMR and LRR. LMR increased and RMR decreased with the increasing N supply. SMR increased with increase N supply up to 80 μmol mol⁻¹ and then leveled off (under elevated [CO₂]) or stated to decline (under ambient [CO₂]) with further increases in N supply. The results suggest that the CO₂ elevation increased biomass accumulation, particularly stem biomass and at higher N supply. The results also suggest that while modest N fertilization will increase seedling growth and biomass accumulation, excessive application of N may not stimulate further growth or even result in growth decline.     

Factors causing variation in fine root biomass in forest ecosystems

Fine roots form one of the most significant components contributing to carbon cycling in forest ecosystems. We study here the effect of variation in root diameter classes, sampling depth and the inclusion of understorey vegetation root biomass in fine root biomass (FRB) estimates. The FRB estimates for different forest biomes are updated using a database of 512 forest stands compiled from the literature. We also investigate the relationships between environmental or forest stand variables and fine root biomass (≤2 mm in diameter) at the stand (g m⁻²) and tree level (g tree⁻¹). The FRB estimates extrapolated for the whole rooting depth were 526 ± 321 g m⁻², 775 ± 474 g m⁻² and 776 ± 518 g m⁻² for boreal, temperate and tropical forests, respectively, and were 26-67% higher than those based on the original sampling depths used. We found significant positive correlations between ≤1 and ≤2 mm diameter roots and between ≤2 and ≤5 mm roots. The FRB estimates, standardized to the ≤2 mm diameter class, were 34-60% higher and 25-29% smaller than those standardized to the ≤1 mm and ≤5 mm diameter classes, respectively. The FRB of the understorey vegetation accounted for 31% of the total FRB in boreal forests and 20% in temperate forests. The results indicate that environmental factors (latitude, mean annual precipitation, elevation, temperature) or forest stand factors (life form, age, basal area, density) can not explain a significant amount of the variation in the total FRB and a maximum of 30% that in the FRB of trees at the stand level, whereas the mean basal area of the forest stand can explain 49% of the total FRB and 79% of the FRB of trees at the tree level.     

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.     

Carbon accumulation along a stand development sequence of Nothofagus antarctica forests across a gradient in site quality in Southern Patagonia

Above- and below-ground C pools were measured in pure even-aged stands of Nothofagusantarctica (Forster f.) Oersted at different ages (5–220 years), crown and site classes in the Patagonian region. Mean tissue C concentration varied from 46.3% in medium sized roots of dominant trees to 56.1% in rotten wood for trees grown in low quality sites. Total C concentration was in the order of: heartwood > rotten wood > sapwood > bark > small branches > coarse roots > leaves > medium roots > fine roots. Sigmoid functions were fitted for total C accumulation and C root/shoot ratio of individual trees against age. Total C accumulated by mature dominant trees was six times greater than suppressed trees in the same stands, and total C accumulated by mature dominant trees grown on the best site quality was doubled that of those on the lowest site quality. Crown classes and site quality also affected the moment of maximum C accumulation, e.g. dominant trees growing on the worse site quality sequestered 0.73 kg C tree⁻¹ year⁻¹ at 139 years compared to the best site where 1.44 kg C tree⁻¹ year⁻¹ at 116 years was sequestered. C root/shoot ratio decreased over time from a maximum value of 1.3–2.2 at 5 years to a steady-state asymptote of 0.3–0.7 beyond 60 years of age depending on site quality. Thus, root C accumulation was greater during the regeneration phase and for trees growing on the poorest sites. The equations developed for individual trees have been used to estimate stand C accumulation from forest inventory data. Total stand C content ranged from 128.0 to 350.9 Mg C ha⁻¹, where the soil C pool represented 52–73% of total ecosystem C depending on age and site quality. Proposed equations can be used for practical purposes such as estimating the impact of silvicultural practices (e.g. thinning or silvopastoral systems) on forest C storage or evaluating the development of both above- and below-ground C over the forest life cycle for different site qualities for accurate quantification of C pools at regional scale.     

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.     

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

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.     

Radiation use efficiency in adjacent hardwood and pine forests in the southern Appalachians

The efficiency with which trees convert photosynthetically active radiation (PAR) to biomass has been shown to be consistent within stands of an individual species, which is useful for estimating biomass production and carbon accumulation. However, radiation use efficiency (?) has rarely been measured in mixed-species forests, and it is unclear how species diversity may affect the consistency of ?, particularly across environmental gradients. We compared aboveground net primary productivity (ANPP), intercepted photosynthetically active solar radiation (IPAR), and radiation use efficiency (? = ANPP/IPAR) between a mixed deciduous forest and a 50-year-old white pine (Pinus strobus L.) plantation in the southern Appalachian Mountains. Average ANPP was similar in the deciduous forest (11.5 Mg ha⁻¹ y⁻¹) and pine plantation (10.2 Mg ha⁻¹ y⁻¹), while ? was significantly greater in the deciduous forest (1.25 g MJ⁻¹) than in the white pine plantation (0.63 g MJ⁻¹). Our results demonstrate that late-secondary hardwood forests can attain similar ANPP as mature P. strobus plantations in the southern Appalachians, despite substantially less annual IPAR and mineral-nitrogen availability, suggesting greater resource-use efficiency and potential for long-term carbon accumulation in biomass. Along a 260 m elevation gradient within each forest there was not significant variation in ?. Radiation use efficiency may be stable for specific forest types across a range of environmental conditions in the southern Appalachian Mountains, and thus useful for generating estimates of ANPP at the scale of individual watersheds.     

Impact of timber harvesting on litterfall inputs and benthic coarse particulate organic matter (CPOM) storage in a small stream draining a eucalyptus plantation

Eucalyptus plantations have a short rotation cycle and harvesting occurs every 12-15 years, with the potential to modify the ecological integrity of the small streams draining the harvested areas through the reduction of litterfall inputs. We studied litterfall inputs and benthic coarse particulate organic matter (CPOM) storage in a small headwater stream draining a eucalyptus (Eucalyptus globulus Labill.) plantation before and after clear felling of the plantation. We hypothesized that wood harvesting will result in a reduction of CPOM inputs and storage in the stream. Litterfall inputs ranged 530-700 g m⁻² y⁻¹ and were approximately halved (200-320 g m⁻² y⁻¹) after the harvesting of the eucalyptus trees. Bark and woody materials showed the largest reduction. Leaf inputs were initially reduced sharply, but, during the second year after the harvest, they recovered to about 90% of the values observed before the harvesting. Harvesting of the eucalyptus plantation caused an increase of benthic CPOM storage to 535 g m⁻², but this was a temporary effect and these materials were washed downstream of the study reach. One year after the harvesting, benthic CPOM was reduced below 15 g m⁻². Bark, twigs and other woody residues generated during the preparation of the logs for transportation were retained within the study site and represented the main component (>90%) of the benthic CPOM after timber harvesting. However, 2 years after the harvesting, low inputs of these materials caused an overall reduction of in-stream retention and residence time of benthic CPOM. Amount and composition of benthic CPOM changed quickly in response to alterations of the riparian forest, so we propose the use of CPOM as an indicator of the impact of forestry activities on the ecological functioning of small streams.     

Responses of oaks and tanoaks to the sudden oak death pathogen after 8 y of monitoring in two coastal California forests

Sudden oak death, caused by Phytophthora ramorum, is widely established in mesic forests of coastal central and northern California. In 2000, we placed 18 plots in two Marin County sites to monitor disease progression in coast live oaks (Quercus agrifolia), California black oaks (Q. kelloggii), and tanoaks (Lithocarpus densiflorus), the species that are most consistently killed by the pathogen in these areas. Through early 2008, the numbers of newly infected trees increased for all species. The infection rate for trees that were asymptomatic in 2000 was 5.0% y⁻¹ for coast live oaks, 4.1% y⁻¹ for black oaks and 10.0% y⁻¹ for tanoaks. Mortality rates were 3.1% y⁻¹ for coast live oaks, 2.4% y⁻¹ for black oaks, and 5.4% y⁻¹ for tanoaks. Mortality not attributed to P. ramorum was 0.54% y⁻¹ for coast live oaks, and 0.75% y⁻¹ for tanoaks. Weibull survival models of trees that were asymptomatic in 2000 provided overall median survival times of 13.7 y for coast live oaks, 13.8 y for black oaks, and 8.8 y for tanoaks. Survival of infected (bleeding) trees declined to 9.7 y for coast live oaks, 6.2 y for black oaks, and 5.8 y for tanoaks. Ambrosia beetle attacks on bleeding trees further reduced modeled survival times by 65–80%, reaffirming the earlier finding that beetle attacks on bleeding cankers considerably reduce survival. Across all plots, the modeled time for 90% of trees that were asymptomatic in 2000 to become infected is 36.5 y for coast live oaks and 15.4 y for tanoaks. There was a trend toward higher infection rates as tree diameter increased. Greater than 90% of living coast live oaks that failed during the study had extensive beetle tunneling at the site of the break. Disease intensity in coast live oaks at the plot level was positively associated with bay laurel (Umbellularia californica) basal area and negatively associated with Pacific madrone (Arbutus menziesii) basal area. This study demonstrates the use of survival modeling to characterize the effects of epidemic disease on different species and to project the future of forests infected with tree pathogens.     

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.     

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.     

Dissolved organic carbon and nitrogen leaching from Scots pine, Norway spruce and silver birch stands in southern Sweden

The effects of three common tree species - Scots pine, Norway spruce and silver birch - on leaching of dissolved organic carbon and dissolved nitrogen were studied in an experimental forest with podzolised soils in southern Sweden. We analyzed soil water collected with lysimeters and modeled water fluxes to estimate dissolved C and N fluxes. Specific UV absorbance (SUVA) was analyzed to get information about the quality of dissolved organic matter leached from the different stands. Under the O horizon, DOC concentrations and fluxes in the birch stands were lower than in the spruce and pine stands; annual fluxes were 21 g m⁻² y⁻¹ for birch and 38 g m⁻² y⁻¹ and 37 g C m⁻² y⁻¹ for spruce and pine, respectively. Under the B horizon, annual fluxes for all tree species ranged between 3 and 5 g C m⁻² y⁻¹, implying greater loss of DOC in the mineral soil in the coniferous stands than in the birch stands. We did not find any effect of tree species on the quality of the dissolved organic matter, as measured by SUVA, indicating that the chemical composition of the organic matter was similar in leachates from all three tree species. Substantial amounts of nitrogen was leached out of the soil profile at the bottom of the B horizon from the pine and birch stands, whereas the spruce stands seemed to retain most of the nitrogen in the soil. These differences in N leaching have implications for soil N budgets.     

Seasonal and spatial variation of nitrogen dynamics in the litter and surface soil layers on a tropical dry evergreen forest slope

Seasonal and spatial variability of litterfall and NO₃⁻ and NH₄⁺ leaching from the litter layer and 5-cm soil depth were investigated along a slope in a tropical dry evergreen forest in northeastern Thailand. Using ion exchange resin and buried bag methods, the vertical flux and transformation of inorganic nitrogen (N) were observed during four periods (dry, early wet, middle wet, and late wet seasons) at 15 subplots in a 180-m × 40-m rectangular plot on the slope. Annual N input via litterfall and inorganic N leached from the litter layer and from 5-cm depth soil were 12.5, 6.9, and 3.7 g N m⁻² year⁻¹, respectively, whereas net mineralization and the inorganic N pool in 0–5-cm soil were 7.1 g N m⁻² year⁻¹ and 1.4 g N m⁻², respectively. During the early wet season (90 days), we observed 82% and 74% of annual NO₃⁻ leaching from the litter layer and 5-cm soil depth, respectively. Higher N input via leaf litterfall in the dry season and via precipitation in the early wet season may have led to higher NO₃⁻ leaching rate from litter and surface soil layers during the early wet season. Large spatial variability in both NO₃⁻ vertical flux and litterfall was also observed within stands. Small-scale spatial patterns of total N input via litterfall were significantly correlated with NO₃⁻ leaching rate from the surface soil layer. In tropical dry evergreen forests, litterfall variability may be crucial to the remarkable seasonal changes and spatial variation in annual NO₃⁻ vertical flux in surface soil layers.     

Wood density in forests of Brazil's ‘arc of deforestation’: Implications for biomass and flux of carbon from land-use change in Amazonia

Wood density is an important variable in estimates of forest biomass and greenhouse-gas emissions from land-use change. The mean wood density used in estimates of forest biomass in the Brazilian Amazon has heretofore been based on samples from outside the “arc of deforestation”, where most of the carbon flux from land-use change takes place. This paper presents new wood density estimates for the southern and southwest Brazilian Amazon (SSWA) portions of the arc of deforestation, using locally collected species weighted by their volume in large local inventories. Mean wood density was computed for the entire bole, including the bark, and taking into account radial and longitudinal variation. A total of 403 trees were sampled at 6 sites. In the southern Brazilian Amazon (SBA), 225 trees (119 species or morpho-species) were sampled at 4 sites. In eastern Acre state 178 trees (128 species or morpho-species) were sampled at breast height in 2 forest types. Mean basic density in the SBA sites was 0.593 ± 0.113 (mean ± 1 S.D.; n = 225; range 0.265–0.825). For the trees sampled in Acre the mean wood density at breast height was 0.540 ± 0.149 (n = 87) in open bamboo-dominated forest and 0.619 ± 0.149 (n = 91) in dense bamboo-free forest. Mean wood density in the SBA sites was significantly higher than in the bamboo dominated forest but not the dense forest at the Acre site. From commercial wood inventories by the RadamBrasil Project in the SSWA portion of the arc of deforestation, the wood volume and wood density of each species or genus were used to estimate average wood density of all wood volume in each vegetation unit. These units were defined by the intersection of mapped forest types and states. The area of each unit was then used to compute a mean wood density of 0.583 g cm⁻³ for all wood volume in the SSWA. This is 13.6% lower than the value applied to this region in previous estimates of mean wood density. When combined with the new estimates for the SSWA, this gave an average wood density of 0.642 g cm⁻³ for all the wood volume in the entire Brazilian Amazon, which is 7% less than a prior estimate of 0.69 g cm⁻³. These results suggest that current estimates of carbon emissions from land-use change in the Brazilian Amazon are too high. The impact on biomass estimates and carbon emissions is substantial because the downward adjustment is greater in forest types undergoing the most deforestation. For 1990, with 13.8 × 10³ km² of deforestation, emissions for the Brazilian Amazon would be reduced by 23.4–24.4 × 10⁶ Mg CO₂-equivalent C/year (for high- and low-trace gas scenarios), or 9.4–9.5% of the gross emission and 10.7% of the net committed emission, both excluding soils.     

Comparison of soil CO2 flux between uncleared and cleared windthrow areas in Estonia and Latvia

Storms can turn a great proportion of forests’ assimilation capacity into dead organic matter because of windthrow and thus its role as a carbon sink will be diminished for some time. However, little is known about the magnitude or extent to which storms affect carbon efflux. We compared soil CO₂ fluxes in wind-thrown forest stands with different time periods since a storm event, and with different management practices (deadwood cleared or left on-site). This study examined changes in soil CO₂ efflux in two windthrow areas in north-eastern Estonia and one area in north-western Latvia, which experienced severe wind storms in the summers of 2001, 2002 and 1967, respectively. We measured soil CO₂ fluxes in stands formerly dominated by Norway spruce (Picea abies L. Karst.) with total and partial canopy destruction (all trees or roughly half of the trees in stand damaged by storm), in harvested areas (material removed after the wind storm) and in control areas (no damage by wind). Removal of wind-damaged material decreased instantaneous CO₂ flux from the soil surface. The highest instantaneous fluxes were measured in areas with total and partial canopy destruction (0.67 g CO₂ m⁻² h⁻¹ in both cases) compared with fluxes in the control areas (0.51 g CO₂ m⁻² h⁻¹), in the new storm-damaged areas where the material was removed (0.57 g CO₂ m⁻² h⁻¹) and in the old storm-damaged area where wood was left on site (0.55 g CO₂ m⁻² h⁻¹). The only factor affecting soil CO₂ flux was location of the measuring collar (plastic collar with diameter 100 mm, height 50 mm) - either on undamaged forest ground or on the uprooted tree pit, where the mineral soil was exposed after disturbance. New wind-thrown stands where residues are left on site would most likely turn to sources of CO₂ for several years until forest regeneration reaches to substantial assimilation rates. New wind-thrown stands where residues are left on site would most likely tend to have elevated CO₂ fluxes for several years until forest regeneration reaches to substantial assimilation rates. However, forest managers might be concerned about the amounts of CO₂ immediately released into the atmosphere if the harvested logs are burned.     

Growth and population structure of the tree species Malouetia tamaquarina (Aubl.) (Apocynaceae) in the central Amazonian floodplain forests and their implication for management

The long-term success of forest management depends primarily on the sustainability of timber production. In this study we analyse the population structure, tree age and wood increment of Malouetia tamaquarina (Aubl.) (Apocynaceae) to define a species-specific minimum logging diameter (MLD) and felling cycle by modelling volume growth. Contrary to other timber species in the nutrient-rich white-water floodplains forests (várzea), M. tamaquarina grows in the subcanopy of old-growth várzea forests. The wood of this species is utilized by local inhabitants in the floodplains for handicraft. In 35 plots of 25 m × 50 m we measured diameter at breast height (DBH) and tree height of all trees taller than 150 cm height. From 37 individuals with DBH > 15 cm we sampled two cores by increment borers to determine the wood density, tree age and diameter increment rates. In the management area of a várzea settlement with about 150 ha recently harvested trees of M. tamaquarina have been recorded and DBH was measured. The species presents an inverse J-shaped diameter distribution indicating that the species is obviously regenerating in the old-growth forests. Tree-ring analysis indicates a mean age of 74.5 years for a DBH of 22.7 cm for a studied population comprising 37 trees with maximum ages of up to 141 years for an individual with a DBH of 45.7 cm. The tree species has low annual diameter increment rates (3.16 ± 0.6 mm) despite a low wood density (0.36 ± 0.05 g cm⁻³). The volume growth model indicates a MLD of 25 cm and a felling cycle of 32.4 years. In the management area 35 trees with a mean DBH of 24 cm were recorded, similar to the defined MLD. The abundance of trees above the MLD is 2.7 trees ha⁻¹, or 405 trees, when extrapolated to the whole management area. Considering a felling cycle of 32.4 years (annual production unit of 4.63 ha) this results in total of 12.5 harvestable trees, almost three times less than actually harvested. The actual practice of harvesting M. tamaquarina risks the overexploitation of this slow-growing species.     

Crown transparency, tree mortality and stem growth of Pinus sylvestris, and colonization of Tomicus piniperda after an outbreak of Gremmeniella abietina

In the year 2000, large areas of forest in Sweden, mainly 30-50 year old Pinus sylvestris (L.) stands, were attacked by the fungus Gremmeniella abietina (Lagerb.) Morelet. The aims of this study were to investigate: (i) the relationship between G. abietina-induced tree crown transparency (CT) and P. sylvestris (L.) tree mortality; (ii) the influence of CT levels on stem growth; (iii) the recovery of the crown; and (iv) the association of CT and colonization by Tomicus piniperda (L.). Thirty-five permanent sample plots were established in five P. sylvestris stands (38-46 years old), infested by G. abietina, and 23 plots in four reference stands, not obviously infested.During the 5 years following the attack, the total mortality amounted to 454 trees ha⁻¹ and 7.8 m² ha⁻¹, on average, in the five infested stands, corresponding to 42% of the trees and 34% of the basal area at the time of the attack. Most of the mortality occurred within 2 years of the attack. The mortality of individual trees (2002-2005) was found to be related to the crown transparency (CT), the position of needle loss within the crown and the tree diameter at breast height. Based on our modeling, the probability of mortality was substantially increased if the initial CT-value was higher than 85%.Growth reductions were detected for individual trees with an initial CT of >c. 40%. In contrast, trees with a low initial CT (<c. 40%) were not affected and even exhibited increased growth. In the five infested stands, the reductions in basal area and volume increment were estimated to be 26-58%, and, 42-73%, respectively, during the five growing seasons after the attacks.The trees in the infested stands that were still alive in spring 2005 had started to recover in terms of CT. Breeding of T. piniperda on the P. sylvestris (L.) stems occurred almost exclusively on stems with a CT > 90%.The data from this study suggest that when a P. sylvestris (L.) stand has been attacked by G. abietina, trees with a CT above 80% should be felled; the remaining trees will have a high probability of survival and resistance to successful breeding by the T. piniperda.