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.     

Variation patterns of fine root biomass,production and turnover in Chinese forests

China’s forests cover 208.3 million ha and span a wide range of climates and a large variety of forest types including tropical,temperate,and boreal forests.However the variation patterns of fine root(2 mm in diameter biomass,production,and turnover from the south to the north are unclear.This study summarizes fine root biomass(FRB),production(FRP)and turnover rate(FRT)in Chi na’s forests as reported by 140 case studies published from 1983 to 2014.The results showed that the mean values o FRB,FRP and FRT in China’s forests were 278 gm~(-2)366 gm~(-2)a~(-1),and 1.19 a~(-1),respectively.Compared with other studies at the regional or global scales,FRB in China’s forests was lower,FRP was similar to estimates a the global scale,but FRT was much higher.FRB,FRP,and FRT in China’s forests increased with increasing mean annual precipitation(MAP),indicating that fine root vari ables were likely related to MAP,rather than mean annua temperature or latitude.This is possibly due to the smal variation in temperature but greater variation in precipitation during the growing season.These findings suggest that spatiotemporal variation in precipitation has a more profound impact on fine root dynamics in China’s forests,and this will impact carbon and nutrient cycles driven by root turnover in the future.     

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.     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

Growth and productivity of black spruce in even- and uneven-aged stands at the limit of the closed boreal forest

The increasing commercial interest and advancing exploitation of new remote territories of the boreal forest require deeper knowledge of the productivity of these ecosystems. Canadian boreal forests are commonly assumed to be evenly aged, but recent studies show that frequent small-scale disturbances can lead to uneven-aged class distributions. However, how age distribution affects tree growth and stand productivity at high latitudes remains an unanswered question. Dynamics of tree growth in even- and uneven-aged stands at the limit of the closed black spruce (Picea mariana) forest in Quebec (Canada) were assessed on 18 plots with ages ranging from 77 to 340 years. Height, diameter and age of all trees were measured. Stem analysis was performed on the 10 dominant trees of each plot by measuring tree-ring widths on discs collected each meter from the stem, and the growth dynamics in height, diameter and volume were estimated according to tree age. Although growth followed a sigmoid pattern with similar shapes and asymptotes in even- and uneven-aged stands, trees in the latter showed curves more flattened and with increases delayed in time. Growth rates in even-aged plots were at least twice those of uneven-aged plots. The vigorous growth rates occurred earlier in trees of even-aged plots with a culmination of the mean annual increment in height, diameter and volume estimated at 40–80 years, 90–110 years earlier than in uneven-aged plots. Stand volume ranged between 30 and 238 m³ ha⁻¹ with 75% of stands showing values lower than 120 m³ ha⁻¹ and higher volumes occurring at greater dominant heights and stand densities. Results demonstrated the different growth dynamics of black spruce in single- and multi-cohort stands and suggested the need for information on the stand structure when estimating the effective or potential growth performance for forest management of this species.     

Transpiration and hydraulic traits of old and regrowth eucalypt forest in southwestern Australia

We tested the hypothesis that overstorey of eucalypt forest dominated by tall, large diameter trees uses less water than regrowth stands in the high rainfall zone (>1100 mm year⁻¹) of the northern jarrah (Eucalyptus marginata) forest in southwestern Australia. We measured leaf area, cover, sapwood area and sapwood density at three paired old and regrowth stands. We also measured sapflow velocity at one paired stand (Dwellingup) from June 2007 to October 2008. Old stands had more basal area but less foliage cover, less leaf area and slightly thinner sapwood. The ratio of sapwood area to basal area decreased markedly as tree size increased. Sapwood area of the regrowth forest stands (6.6 ± 0.30 m² ha⁻¹) was nearly double that of the old stands (3.4 ± 0.17 m² ha⁻¹), despite larger basal area at the old stands. Leaf area index of the regrowth stands (2.1 ± 0.26) was only one-third larger than that at the old stands (1.5 ± 0.15); hence, the ratio of leaf area to sapwood area was larger in old stands than in regrowth stands (0.45 ± 0.022 m² cm⁻² versus 0.32 ± 0.045 m² cm⁻²). Our results are consistent with theories that trees have evolved to optimize carbon gain rather than maintain stomatal conductance. Neither sapwood density (540–650 kg m⁻³) nor sap velocity differed greatly between regrowth and old stands. At the old forest site, daily transpiration rose from 0.5 mm day⁻¹ in winter to 0.9 mm day⁻¹ in spring–summer, compared to 0.9 mm day⁻¹ and 1.8 mm day⁻¹ at the regrowth site. Annual water use by the overstorey trees was estimated to be ∼230 mm year⁻¹ for the old stand and ∼500 mm year⁻¹ at the regrowth stand, or 20% and 44% of annual rainfall. The overwhelming role of stand sapwood area in determining stand water use, combined with the marked changes in the ratio of sapwood area to basal area with tree age and size, suggest that stand overstorey structure can be managed to alter overstorey water use and catchment water yield. Silviculture to promote old-forest-like attributes may be a viable means of delivering multiple water and conservation benefits.     

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.     

Profound impoverishment of the large-tree stand in a hyper-fragmented landscape of the Atlantic forest

Large tree species have a disproportional influence on the structure and functioning of tropical forests, but the forces affecting their long-term persistence in human-dominated landscapes remain poorly understood. Here we test the hypothesis that aging forest edges and small fragments (3.4–295.7 ha) are greatly impoverished in terms of species richness and abundance of large trees in comparison to core areas of forest interior. The study was conducted in a hyper-fragmented landscape of the Atlantic forest, northeast Brazil. Large tree species were quantified by recording all trees (DBH ≥ 10 cm) within fifty-eight 0.1-ha plots distributed in three forest habitats: small forest fragments (n = 28), forest edges (n = 10), and primary forest interior areas within an exceptional large forest remnant (n = 20). Large tree species and their stems ≥10 cm DBH were reduced by half in forest edges and fragments. Moreover, these edge-affected habitats almost lacked large-stemmed trees altogether (0.24 ± 0.27% of all stems sampled), and very tall trees were completely absent from forest edges. In contrast, large trees contributed to over 1.5% of the whole stand in forest interior plots (2.9 ± 2.8%). Habitats also differed in terms of tree architecture: relative to their DBH trees were on average 30% shorter in small fragments and forest edges. Finally, an indicator species analysis yielded an ecological group of 12 large tree species that were significantly associated with forest interior plots, but were completely missing from edge-affected habitats. Our results suggest a persistent and substantial impoverishment of the large-tree stand, including the structural collapse of forest emergent layer, in aging, hyper-fragmented landscapes.     

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.     

Dominance by the introduced tree Rhamnus cathartica (common buckthorn) may limit aboveground carbon storage in Southern Wisconsin forests

Many ecosystems are now dominated by introduced species, and because dominant species drive ecosystem properties, these changes lead to increased uncertainty in estimates of carbon storage and cycling. We examined aboveground biomass in forests dominated by the introduced tree Rhamnus cathartica (common buckthorn) relative to forests dominated by native species, and measured aboveground biomass increment over a three-year period (2005-2008). Three of the four lowest biomass levels occurred in R. cathartica-dominated forests, and biomass in these forest types was stored primarily in trees 10-20 cm DBH. By contrast, forests dominated by native trees (including those with R. cathartica understories) had the six highest biomass levels, and biomass was stored primarily in trees >50 cm DBH. On average, forests dominated by R. cathartica stored half as much aboveground biomass (14.6 ± 3.3 kg/m²) as forests dominated by native tree species (28.9 ± 8.3 kg/m²). R. cathartica-dominated forests also had half the aboveground biomass increment of native-dominated forests (0.28 vs. 0.60 kg/m²/year). Although known anecdotally as a fast-growing species, R. cathartica growth rates declined with increasing size. Between 2005 and 2008, R. cathartica individuals <10 cm DBH grew faster than native species; however, R. cathartica individuals >10 cm DBH grew consistently slower than native species. Overall, our findings indicate that intrinsic size limitations on R. cathartica will lead to lower biomass stocks in forests where it acts as a canopy dominant relative to forests dominated by native tree species.     

Comparing aboveground carbon sequestration between moso bamboo (Phyllostachys heterocycla) and China fir (Cunninghamia lanceolata) forests based on the allometric model

The purpose of this study was to compare carbon sequestration between moso bamboo (Phyllostachys heterocycla) and China fir (Cunninghamia lanceolata) forests. The study site was located in the lower mountain area of central Taiwan, where both moso bamboo and China fir were rich. In addition, moso bamboo and China fir forests were surveyed on 12 and 19 plantations, respectively. We predicted carbon sequestration based on the allometric model for moso bamboo and China fir forests and compared the relationships between characteristics of bamboo forests and elevation. The results showed that mean diameter at breast height (DBH), culms per hectare and aboveground biomass were not clearly affected by elevation, whereas a negative correlation (R = −0.600, p = 0.039) between mean DBH and stand density was found for moso bamboo forests. Moreover, the aboveground carbon storage was higher for China fir forests than for moso bamboo (99.5 vs. 40.6 Mg ha⁻¹). However, moso bamboo is an uneven-aged stand which is only composed of 1-5-year-old culms, while China fir is an even-aged stand and the age range is from 15 to 54 years, such that, per year, the mean aboveground carbon sequestration is 8.13 ± 2.15 and 3.35 ± 2.02 Mg ha⁻¹ for moso bamboo and China fir, respectively. On the other hand, the mean carbon sequestration of China fir decreases with increasing the age class. Furthermore, the ratio of moso bamboo to China fir is 2.39 and a T-test showed that the aboveground carbon levels were significantly different between these two species; thus, moso bamboo is a species with high potential for carbon sequestration.     

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.     

Midcanopy growth following thinning in young-growth conifer forests on the Olympic Peninsula western Washington

Midcanopy layers are essential structures in “old-growth” forests on the Olympic Peninsula. Little is known about which stand and tree factors influence the ability of midcanopy trees in young-growth forests to respond to release; however, this information is important to managers interested in accelerating development of late-successional structural characteristics. We examined basal area growth response of midcanopy trees following variable-density thinning in an effort to determine the effect of thinning and local environment on the release of western hemlock (Tsuga heterophylla (Raf.) Sarg.) and western redcedar (Thuja plicata ex. D. Don) on the Olympic Peninsula in western Washington. Release was measured as the difference between average annual basal area growth over the 5-year prior to thinning and the 3-to-6 year period following thinning. Results indicate that while growth rates were similar prior to thinning (5.4 cm² year⁻¹in both thinned and unthinned patches) midcanopy trees retained in a uniformly thinned matrix grew significantly more (8.0 cm² year⁻¹) than those in unthinned patches (5.4 cm² year⁻¹) for western hemlock and for western redcedar. Crown fullness and crown crowding affected the release of western hemlock in the thinned matrix. Initial tree size, relative age, local crowding and measures of crown size and vigor affected the release of western redcedar in the thinned matrix. Our results indicate that midcanopy western hemlock and western redcedar retain the ability to respond rapidly with increased growth when overstory competition is reduced and the magnitude of response is related to neighborhood variables (intracohort competition, overstory competition, and tree vigor), thus suggest that variable-density thinning can be an effective tool to create variability in the growth of midcanopy trees in young-growth stands. We expect that this rapid response will produce even greater variability over time.     

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.     

Regeneration of Populus nine years after variable retention harvest in boreal mixedwood forests

Aspen and balsam poplar regeneration from root suckers were assessed in boreal mixedwood forests nine years after logging in a variable retention experiment (EMEND Project—Ecosystem Management Emulating Natural Disturbance) located north of Peace River, Alberta, Canada. Five levels of retention of mature trees (2%, 10%, 20%, 50% or 75% of the original basal area) were applied in stands dominated by aspen, white spruce or mixtures of the two species. Basal area of aspen (or that of aspen plus balsam poplar combined) prior to logging strongly influenced sucker density of aspen (or aspen + balsam poplar combined) and in some cases their growth. Nine years after harvest there was a decline in sucker density and volume ha⁻¹ with increasing retention levels of aspen (or both poplars combined); sucker density declined by 50% when only 20% of the original basal area was left in the stand. Retaining mature spruce trees in the stand had little influence on the number of suckers but did affect their total volume ha⁻¹. Thus, we suggest that by knowing stand aspen and balsam poplar density prior to logging and varying levels of retention of aspen and balsam poplar or conifers at harvest, the density of Populus regeneration can be predicted by managers, thereby allowing them to create a range of mixedwood conditions.     

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.     

Soil–atmosphere CO2, CH4 and N2O fluxes in boreal forestry-drained peatlands

Greenhouse gas emissions from managed peatlands are annually reported to the UNFCCC. For the estimation of greenhouse gas (GHG) balances on a country-wide basis, it is necessary to know how soil–atmosphere fluxes are associated with variables that are available for spatial upscaling. We measured momentary soil–atmosphere CO₂ (heterotrophic and total soil respiration), CH₄ and N₂O fluxes at 68 forestry-drained peatland sites in Finland over two growing seasons. We estimated annual CO₂ effluxes for the sites using site-specific temperature regressions and simulations in half-hourly time steps. Annual CH₄ and N₂O fluxes were interpolated from the measurements. We then tested how well climate and site variables derived from forest inventory results and weather statistics could be used to explain between-site variation in the annual fluxes. The estimated annual CO₂ effluxes ranged from 1165 to 4437 g m⁻² year⁻¹ (total soil respiration) and from 534 to 2455 g m⁻² year⁻¹ (heterotrophic soil respiration). Means of 95% confidence intervals were ±12% of total and ±22% of heterotrophic soil respiration. Estimated annual CO₂ efflux was strongly correlated with soil respiration at the reference temperature (10 °C) and with summer mean air temperature. Temperature sensitivity had little effect on the estimated annual fluxes. Models with tree stand stem volume, site type and summer mean air temperature as independent variables explained 56% of total and 57% of heterotrophic annual CO₂ effluxes. Adding summer mean water table depth to the models raised the explanatory power to 66% and 64% respectively. Most of the sites were small CH₄ sinks and N₂O sources. The interpolated annual CH₄ flux (range: −0.97 to 12.50 g m⁻² year⁻¹) was best explained by summer mean water table depth (r² = 64%) and rather weakly by tree stand stem volume (r² = 22%) and mire vegetation cover (r² = 15%). N₂O flux (range: −0.03 to 0.92 g m⁻² year⁻¹) was best explained by peat CN ratio (r² = 35%). Site type explained 13% of annual N₂O flux. We suggest that water table depth should be measured in national land-use inventories for improving the estimation of country-level GHG fluxes for peatlands.     

Allometric equations for tree species and carbon stocks for forests of northwestern Mexico

Allometric equations were developed and applied to forest inventory data to estimate biomass and carbon stocks for temperate species and forests of Durango and Chihuahua and for tropical dry forests of Sinaloa, Mexico. A total of 872 trees were harvested and dissected into their component parts: leaves and branches, boles, and coarse roots. Coarse roots of 40 temperate trees ranging in diameter at breast height (DBH) from 6.0 to 52.9 cm were excavated in their entirety (i.e., >0.5 cm diameter). The species sampled (number of trees) in tropical dry forests (39) were Lysiloma divaricata (Jacq) Macbr. (10), Haematoxylon brasiletto Karst. (10), Cochlospermum vitifolium (Wild.) (5), Ceiba acuminata (S. Watson) Rose (5), Bursera penicillata (B. inopinnata) (5), and Jatropha angustifolia Mull. Arg. (4) and in temperate forests (833) were Quercus spp. (118) (Q. rugosa Neé, 15, Quercus sideroxylla Humb. & Bonpl, 51, Quercus spp., 52), Pinus herrerae Martinez 1940 (19), Pinus oocarpa Schiede ex Schlectendal 1838 (31), Pinus engelmannii Carriere 1854 (7), Psudotsuga menziesii (Mirb.) Franco (19), Pinus leiophylla Schiede ex Schlectendal et Chamisso 1831 (27), Pinus teocote Schiede ex Schlectendal et Chamisso (55), Pinus ayacahuite Ehrenb. ex Schltdl. (58), Pinus cooperi Blanco (48), Pinus durangensis Martinez 1942 (385), and Pinus arizonica Engelmann 1879 (66). Allometric equations having only DBH as an independent variable were developed for each component of each species. Since Pinus herrerae, Pinus engelmannii, Pinus oocarpa and Pseudotsuga menziensii had a small number of trees, an individual allometric equation was developed for these species. We used non-linear regression to fit parameters of the typical allometric power equation. The resulting 31 equations (10 species or groups of species, three biomass components; bole, branch and leaves, and total aerial; and the generalized equation for coarse roots) fit the data well and enable the user to predict biomass by component for each of the 10 different groups of species or each of six temperate species. A single allometric equation that incorporates the basic specific gravity for aboveground biomass of all temperate tree species also fit the data well, and this equation provides both the detail and the accuracy supplied by species-specific, plant-part-specific equations. Biomass equations coupled with forest inventory data for temperate (637 circular, 1/10 ha plots) and tropical dry forests (166 20 m × 20 m-quadrats) of northwestern Mexico predict a mean (confidence intervals) of 130 Mg ha⁻¹ (4.2 Mg ha⁻¹) and 73 Mg ha⁻¹ (7.1 Mg ha⁻¹) for total tree and total aboveground biomass, respectively. Large sample sizes and the economic and ecological importance of the species studied make this data set uniquely useful for biomass estimations and for understanding the inherent heterogeneity of tree structure in dynamic tropical and temperate environments of northwestern Mexico.     

Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Tree height in Brazil's ‘arc of deforestation’: Shorter trees in south and southwest Amazonia imply lower biomass

This paper estimates the difference in stand biomass due to shorter and lighter trees in southwest (SW) and southern Amazonia (SA) compared to trees in dense forests in central Amazonia (CA). Forest biomass values used to estimate carbon emissions from deforestation throughout, Brazilian Amazonia will be affected by any differences between CA forests and those in the “arc of deforestation” where clearing activity is concentrated along the southern edge of the Amazon forest. At 12 sites (in the Brazilian states of Amazonas, Acre, Mato Grosso and Pará) 763 trees were felled and measurements were made of total height and of stem diameter. In CA dense forest, trees are taller at any given diameter than those in SW bamboo-dominated open, SW bamboo-free dense forest and SA open forests. Compared to CA, the three forest types in the arc of deforestation occur on more fertile soils, experience a longer dry season and/or are disturbed by climbing bamboos that cause frequent crown damage. Observed relationships between diameter and height were consistent with the argument that allometric scaling exponents vary in forests on different substrates or with different levels of natural disturbance. Using biomass equations based only on diameter, the reductions in stand biomass due to shorter tree height alone were 11.0, 6.2 and 3.6%, respectively, in the three forest types in the arc of deforestation. A prior study had shown these forest types to have less dense wood than CA dense forest. When tree height and wood density effects were considered jointly, total downward corrections to estimates of stand biomass were 39, 22 and 16%, respectively. Downward corrections to biomass in these forests were 76 Mg ha⁻¹ (∼21.5 Mg ha⁻¹ from the height effect alone), 65 Mg ha⁻¹ (18.5 Mg ha⁻¹ from height), and 45 Mg. ha⁻¹ (10.3 Mg ha⁻¹ from height). Hence, biomass stock and carbon emissions are overestimated when allometric relationships from dense forest are applied to SW or SA forest types. Biomass and emissions estimates in Brazil's National Communication under the United Nations Framework Convention on Climate Change require downward corrections for both wood density and tree height.