Elevation effects on the carbon budget of tropical mountain forests (S Ecuador): the role of the belowground compartment

Carbon storage and sequestration in tropical mountain forests and their dependence on elevation and temperature are not well understood. In an altitudinal transect study in the South Ecuadorian Andes, we tested the hypotheses that (i) aboveground net primary production (ANPP) decreases continuously with elevation due to decreasing temperatures, whereas (ii) belowground productivity (BNPP) remains constant or even increases with elevation due to a shift from light to nutrient limitation of tree growth. In five tropical mountain forests between 1050 and 3060 m a.s.l., we investigated all major above‐ and belowground biomass and productivity components, and the stocks of soil organic carbon (SOC). Leaf biomass, stemwood mass and total aboveground biomass (AGB) decreased by 50% to 70%, ANPP by about 70% between 1050 and 3060 m, while stem wood production decreased 20‐fold. Coarse and large root biomass increased slightly, fine root biomass fourfold, while fine root production (minirhizotron study) roughly doubled between 1050 and 3060 m. The total tree biomass (above‐ and belowground) decreased from about 320 to 175 Mg dry mass ha^?1, total NPP from ca. 13.0 to 8.2 Mg ha^?1 yr^?1. The belowground/aboveground ratio of biomass and productivity increased with elevation indicating a shift from light to nutrient limitation of tree growth. We propose that, with increasing elevation, an increasing nitrogen limitation combined with decreasing temperatures causes a large reduction in stand leaf area resulting in a substantial reduction of canopy carbon gain toward the alpine tree line. We conclude that the marked decrease in tree height, AGB and ANPP with elevation in these mountain forests is caused by both a belowground shift of C allocation and a reduction in C source strength, while a temperature‐induced reduction in C sink strength (lowered meristematic activity) seems to be of secondary importance.     

Estimating Fine Root Turnover in Tropical Forests along an Elevational Transect using Minirhizotrons

Growth and death of fine roots represent an important carbon sink in forests. Our understanding of the patterns of fine root turnover is limited, in particular in tropical forests, despite its acknowledged importance in the global carbon cycle. We used the minirhizotron technique for studying the changes in fine root longevity and turnover along a 2000-m-elevational transect in the tropical mountain forests of South Ecuador. Fine root growth and loss rates were monitored during a 5-mo period at intervals of four weeks with each 10 minirhizotron tubes in three stands at 1050, 1890, and 3060 m asl. Average root loss rate decreased from 1.07 to 0.72 g/g/yr from 1050 to 1890 m, indicating an increase in mean root longevity with increasing elevation. However average root loss rate increased again toward the uppermost stand at 3060 m (1.30 g/g/yr). Thus, root longevity increased from lower montane to mid-montane elevation as would be expected from an effect of low temperature on root turnover, but it decreased further upslope despite colder temperatures. We suggest that adverse soil conditions may reduce root longevity at high elevations in South Ecuador, and are thus additional factors besides temperature that control root dynamics in tropical mountain forests.     

Forest aboveground biomass along an elevational transect in Sulawesi,Indonesia, and the role of Fagaceae in tropical montane rain forests

Aim This study investigates how estimated tree aboveground biomass (AGB) of tropical montane rain forests varies with elevation, and how this variation is related to elevational change in floristic composition, phylogenetic community structure and the biogeography of the dominant tree taxa. Location Lore Lindu National Park, Sulawesi, Indonesia. Methods Floristic inventories and stand structural analyses were conducted on 13 plots (each 0.24 ha) in four old‐growth forest stands at 1050, 1400, 1800 and 2400 m a.s.l. (submontane to upper montane elevations). Tree AGB estimates were based on d.b.h., height and wood specific gravity. Phylogenetic diversity and biogeographical patterns were analysed based on tree family composition weighted by AGB. Elevational trends in AGB were compared with other Southeast Asian and Neotropical transect studies (n = 7). Results AGB was invariant from sub‐ to mid‐montane elevation (309–301 Mg ha^?1) and increased slightly to 323 Mg ha^?1 at upper montane elevation. While tree and canopy height decreased, wood specific gravity increased. Magnoliids accounted for most of the AGB at submontane elevations, while eurosids I (including Fagaceae) contributed substantially to AGB at all elevations. Phylogenetic diversity was highest at upper montane elevations, with co‐dominance of tree ferns, Podocarpaceae, Trimeniaceae and asterids/euasterids II, and was lowest at lower/mid‐montane elevations, where Fagaceae contributed > 50% of AGB. Biogeographical patterns showed a progression from dominant tropical families at submontane to tropical Fagaceae (Castanopsis, Lithocarpus) at lower/mid‐montane, and to conifers and Australasian endemics at upper montane elevations. Cross‐continental comparisons revealed an elevational AGB decrease in transects with low/no presence of Fagaceae, but relatively high AGB in montane forests with moderate to high abundance of this family. Main conclusions AGB is determined by both changes in forest structure and shifts in species composition. In our study, these two factors traded off so that there was no net change in AGB, even though there were large changes in forest structure and composition along the elevational gradient. Southeast Asian montane rain forests dominated by Fagaceae constitute important carbon stocks. The importance of biogeography and species traits for biomass estimation should be considered by initiatives to reduce emissions from deforestation and forest degradation (REDD) and in taxon choice in reforestation for carbon offsetting.     

Below-ground response of Norway spruce to climate conditions at Mt. Brocken (Germany)—A re-assessment of Central Europe's northernmost treeline

Alpine treelines at medium high mountains are less abundant and thus have been less frequently studied than at high-elevation mountain ranges of the world. We studied mature Norway spruce stands along an extended elevation transect at Mt. Brocken (Harz Mountains, Central Germany) to analyse the altitudinal changes in climate-related growth conditions, and to evaluate the prevailing climate conditions at the treeline of medium high Mt. Brocken. A particular aim was to analyse the change in fine root biomass partitioning along the transect towards the treeline. Microclimate conditions at the treeline of Mt. Brocken were very similar to other treeline sites worldwide. Tree height and stem biomass strongly decreased from middle elevations towards the treeline. On the contrary, fine root biomass and the ratio of fine root/stem biomass strongly increased towards the treeline indicating a marked shift in carbon allocation in favour of the fine root system with elevation. A meta-analysis of literature data revealed that the elevation-related increase in dry mass partitioning to the fine root system is a general phenomenon for Norway spruce stands in northern and central European mountains. We conclude that the particularly large fine root system of Norway spruce at cold sites represents a mechanism to cope with unfavourable soil conditions such as reduced or temporally variable nutrient supply.     

Optimal climate for large trees at high elevations drives patterns of biomass in remote forests of Papua New Guinea

Our ability to model global carbon fluxes depends on understanding how terrestrial carbon stocks respond to varying environmental conditions. Tropical forests contain the bulk of the biosphere's carbon. However, there is a lack of consensus as to how gradients in environmental conditions affect tropical forest carbon. Papua New Guinea (PNG) lies within one of the largest areas of contiguous tropical forest and is characterized by environmental gradients driven by altitude; yet, the region has been grossly understudied. Here, we present the first field assessment of aboveground biomass (AGB) across three main forest types of PNG using 193 plots stratified across 3,100‐m elevation gradient. Unexpectedly, AGB had no direct relationship to rainfall, temperature, soil, or topography. Instead, natural disturbances explained most variation in AGB. While large trees (diameter at breast height > 50 cm) drove altitudinal patterns of AGB, resulting in a major peak in AGB (2,200–3,100 m) and some of the most carbon‐rich forests at these altitudes anywhere. Large trees were correlated to a set of climatic variables following a hump‐shaped curve. The set of “optimal” climatic conditions found in montane cloud forests is similar to that of maritime temperate areas that harbor the largest trees in the world: high ratio of precipitation to evapotranspiration (2.8), moderate mean annual temperature (13.7°C), and low intra‐annual temperature range (7.5°C). At extreme altitudes (2,800–3,100 m), where tree diversity elsewhere is usually low and large trees are generally rare or absent, specimens from 18 families had girths >70 cm diameter and maximum heights 20–41 m. These findings indicate that simple AGB‐climate‐edaphic models may not be suitable for estimating carbon storage in forests where optimal climate niches exist. Our study, conducted in a very remote area, suggests that tropical montane forests may contain greater AGB than previously thought and the importance of securing their future under a changing climate is therefore enhanced.     

Altitudinal Change in LAI and Stand Leaf Biomass in Tropical Montane Forests: a Transect Study in Ecuador and a Pan-Tropical Meta-Analysis

Abstract Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm² g⁻¹ whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.     

Airborne LiDAR technique for estimating biomass components of maize: A case study in Zhangye City,Northwest China

Crop biomass is an important ecological indicator of growth, light use efficiency, and carbon stocks in agro-ecosystems. Light detection and ranging (LiDAR) or laser scanning has been widely used to estimate forest structural parameters and biomass. However, LiDAR is rarely used to estimate crop parameters because the short, dense canopies of crops limit the accuracy of the results. The objective of this study is to explore the potential of airborne LiDAR data in estimating biomass components of maize, namely aboveground biomass (AGB) and belowground biomass (BGB). Five biomass-related factors were measured during the entire growing season of maize. The field-measured canopy height and leaf area index (LAI) were identified as the factors that most directly affect biomass components through Pearson's correlation analysis and structural equation modeling (SEM). Field-based estimation models were proposed to estimate maize biomass components during the tasseling stage. Subsequently, the maize height and LAI over the entire study area were derived from LiDAR data and were used as input for the estimation models to map the spatial pattern of the biomass components. The results showed that the LiDAR-estimated biomass was comparable to the field-measured biomass, with root mean squared errors (RMSE) of 288.51 g/m² (AGB), and 75.81 g/m² (BGB). In conclusion, airborne LiDAR has great potential for estimating canopy height, LAI, and biomass components of maize during the peak growing season.     

Above- and belowground biomass in relation to environmental factors in temperate grasslands, Inner Mongolia

Above- and belowground biomasses of grasslands are important parameters for characterizing re- gional and global carbon cycles in grassland ecosystems. Compared with the relatively detailed in- formation for aboveground biomass (AGB), belowground biomass (BGB) is poorly reported at the re- gional scales. The present study, based on a total of 113 sampling sites in temperate grassland of the Inner Mongolia, investigated regional distribution patterns of AGB, BGB, vertical distribution of roots, and their relationships with environmental factors. AGB and BGB increased from the southwest to the northeast of the study region. The largest biomass occurred in meadow steppe, with mean AGB and BGB of 196.7 and 1385.2 g/m2, respectively; while the lowest biomass occurred in desert steppe, with an AGB of 56.6 g/m2 and a BGB of 301.0 g/m2. In addition, about 47% of root biomass was distributed in the top 10 cm soil. Further statistical analysis indicated that precipitation was the primary determinant factor in shaping these distribution patterns. Vertical distribution of roots was significantly affected by precipitation, while the effects of soil texture and grassland types were weak.     

Fine root biomass under light gap openings in an Amazon rain forest

Summary Belowground processes in light gap openings are poorly understood, particularly in tropical forests. Fine roots in three zones of light gap openings and adjacent intact forest were regularly measured in buried bags and surface litter envelopes for 2 years. Fine root biomass does not vary significantly within gaps for either buried bags or for surface litter envelopes. When entire gaps are compared without regard for within gap zones, root growth into both surface litter and buried bags is significantly different between gaps, with highest rates of fine root biomass accumulation in the smallest gap. These results suggest that the aboveground within-gap zones do not result in a congruent pattern of below-ground zonation. Gap size, decomposition of the fallen tree, and pre-gap fine root growth rates should be considered to determine fine root growth patterns following the formation of light gap openings.     

Pattern and control of biomass allocation across global forest ecosystems

The underground part of a tree is an important carbon sink in forest ecosystems. Understanding biomass allocation between the below‐ and aboveground parts (root:shoot ratios) is necessary for estimation of the underground biomass and carbon pool. Nevertheless, large‐scale biomass allocation patterns and their control mechanisms are not well identified. In this study, a large database of global forests at the community level was compiled to investigate the root:shoot ratios and their responses to environmental factors. The results indicated that both the aboveground biomass (AGB) and belowground biomass (BGB) of the forests in China (medians 73.0 Mg/ha and 17.0 Mg/ha, respectively) were lower than those worldwide (medians 120.3 Mg/ha and 27.7 Mg/ha, respectively). The root:shoot ratios of the forests in China (median = 0.23), however, were not significantly different from other forests worldwide (median = 0.24). In general, the allocation of biomass between the belowground and aboveground parts was determined mainly by the inherent allometry of the plant but also by environmental factors. In this study, most correlations between root:shoot ratios and environmental factors (development parameter, climate, altitude, and soil) were weak but significant (p < .01). The allometric model agreed with the trends observed in this study and effectively estimated BGB based on AGB across the entire database.     

Effects of soil C:N:P stoichiometry on biomass allocation in the alpine and arid steppe systems

Soil nutrients strongly influence biomass allocation. However, few studies have examined patterns induced by soil C:N:P stoichiometry in alpine and arid ecosystems. Samples were collected from 44 sites with similar elevation along the 220‐km transect at spatial intervals of 5 km along the northern Tibetan Plateau. Aboveground biomass (AGB) levels were measured by cutting a sward in each plot. Belowground biomass (BGB) levels were collected from soil pits in a block of 1 m × 1 m in actual root depth. We observed significant decreases in AGB and BGB levels but increases in the BGB:AGB ratio with increases in latitude. Although soil is characterized by structural complexity and spatial heterogeneity, we observed remarkably consistent C:N:P ratios within the cryic aridisols. We observed significant nonlinear relationships between the soil N:P and BGB:AGB ratios. The critical N:P ratio in soils was measured at approximately 2.0, above which the probability of BGB:AGB response to nutrient availability is small. These findings serve as interesting contributions to the global data pool on arid plant stoichiometry, given the previously limited knowledge regarding high‐altitude regions.     

Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above‐ and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha^?1 at 194 m to 4.95 ± 0.62 Mg C ha^?1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha^?1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha^?1 at 194 m to 47.03 Mg C ha^?1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha^?1 yr^?1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha^?1 yr^?1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha^?1 yr^?1 (210 m) and 2.59 ± 0.40 Mg C ha^?1 yr^?1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above‐ and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.     

Changes in individual plant traits and biomass allocation in alpine meadow with elevation variation on the Qinghai-Tibetan Plateau

Plant traits and individual plant biomass allocation of 57 perennial herbaceous species, belonging to three common functional groups (forbs, grasses and sedges) at subalpine (3700 m ASL), alpine (4300 m ASL) and subnival (⩾5000 m ASL) sites were examined to test the hypothesis that at high altitudes, plants reduce the proportion of aboveground parts and allocate more biomass to belowground parts, especially storage organs, as altitude increases, so as to geminate and resist environmental stress. However, results indicate that some divergence in biomass allocation exists among organs. With increasing altitude, the mean fractions of total biomass allocated to aboveground parts decreased. The mean fractions of total biomass allocation to storage organs at the subalpine site (7%±2% S.E.) were distinct from those at the alpine (23%±6%) and subnival (21%±6%) sites, while the proportions of green leaves at all altitudes remained almost constant. At 4300 m and 5000 m, the mean fractions of flower stems decreased by 45% and 41%, respectively, while fine roots increased by 86% and 102%, respectively. Specific leaf areas and leaf areas of forbs and grasses deceased with rising elevation, while sedges showed opposite trends. For all three functional groups, leaf area ratio and leaf area root mass ratio decreased, while fine root biomass increased at higher altitudes. Biomass allocation patterns of alpine plants were characterized by a reduction in aboveground reproductive organs and enlargement of fine roots, while the proportion of leaves remained stable. It was beneficial for high altitude plants to compensate carbon gain and nutrient uptake under low temperature and limited nutrients by stabilizing biomass investment to photosynthetic structures and increasing the absorption surface area of fine roots. In contrast to forbs and grasses that had high mycorrhizal infection, sedges had higher single leaf area and more root fraction, especially fine roots.     

Stocks of carbon and nitrogen and partitioning between above- and belowground pools in the Brazilian coastal Atlantic Forest elevation range

We estimated carbon and nitrogen stocks in aboveground biomass (AGB) and belowground biomass (BGB) along an elevation range in forest sites located on the steep slopes of the Serra do Mar on the north coast of the State of São Paulo, southeast Brazil. In elevations of 100 m (lowland), 400 m (submontane), and 1000 m (montane) four 1-ha plots were established, and above- (live and dead) and belowground (live and dead) biomass were determined. Carbon and nitrogen concentrations in each compartment were determined and used to convert biomass into carbon and nitrogen stocks. The carbon aboveground stock (C_AGB) varied along the elevation range from approximately 110 to 150 Mg·ha⁻¹, and nitrogen aboveground stock (N_AGB), varied from approximately 1.0 to 1.9 Mg·ha⁻¹. The carbon belowground stock (C_BGB) and the nitrogen belowground stock (N_BGB) were significantly higher than the AGB and varied along the elevation range from approximately 200–300 Mg·ha⁻¹, and from 14 to 20 Mg·ha⁻¹, respectively. Finally, the total carbon stock (C_TOTAL) varied from approximately 320 to 460 Mg·ha⁻¹, and the nitrogen total stock (N_TOTAL) from approximately 15 to 22 Mg·ha⁻¹. Most of the carbon and nitrogen stocks were found belowground and not aboveground as normally found in lowland tropical forests. The above- and belowground stocks, and consequently, the total stocks of carbon and nitrogen increased significantly with elevation. As the soil and air temperature also decreased significantly with elevation, we found a significantly inverse relationship between carbon and nitrogen stocks and temperature. Using this inverse relationship, we made a first approach estimate that an increase of 1°C in soil temperature would decrease the carbon and nitrogen stocks in approximately 17 Mg·ha⁻¹ and 1 Mg·ha⁻¹ of carbon and nitrogen, respectively.     

Root functioning in tropical high-elevation forests: Environmental vs. biological control of root water absorption

Lowered temperatures may reduce the root water uptake of tropical trees at high elevations through several mechanisms; however, field studies to test their relevance are lacking. We measured sap flux density (J) in small-diameter tree roots across a 2000-m elevation transect in a tropical mountain forest for quantifying the effects of temperature (T_a), VPD and soil moisture (θ) on root water flow and uptake at different elevations. Recently developed miniature heat balance-sap flow gauges were applied to roots of about 10 mm in diameter in mountain forest stands at 1050, 1890 and 3060 m a.s.l. in the Ecuadorian Andes and the measured flow was related to anatomical properties of the root xylem. Between 1050 and 3060 m, mean J decreased to about a third. VPD was the most influential environmental factor controlling J at 1050 and 1890 m, while T_a was the key determinant at 3060 m. Large vessels were absent in the root xylem of high-elevation trees which resulted in a 10-fold decrease of theoretical hydraulic conductivity (k_h^theor) between 1050 and 3060 m. We conclude that both physical limitations (reduced VPD, increased viscosity of water) and biological constraints (large decrease of k_h^theor) result in a significantly reduced J and root water uptake of the trees in high-elevation tropical forests.     

Tropical moist <Emphasis Type="Italic">Polylepis</Emphasis> stands at the treeline in East Bolivia: the effect of elevation on stand microclimate,above- and below-ground structure,and regeneration

We studied Polylepis forests along an elevational transect between 3,650 and 4,050 m a.s.l. at the treeline of the moist eastern cordillera in Bolivia to examine changes in above- and below-ground stand structure, leaf and root morphology, and regeneration in relation to stand microclimate. Field measurements and model predictions indicated relatively cold growth conditions of the Polylepis forests. Tree height, stem diameter, and basal area of the stands decreased markedly while stem density increased with elevation. Leaf morphology differed between the two occurring Polylepis species, and trees at the treeline had smaller leaves with higher specific leaf area. In contrast, fine root biomass increased from 37 g m⁻² at the lowermost stand to 234 g m⁻² at the treeline. Trees of the uppermost stand had higher specific root surface area and a much higher number of root tips per unit dry mass. Thus, root surface area and total number of root tips per unit ground area increased conspicuously from the lowermost stand to the treeline. Density of young growth inside the forest increased towards the treeline, while density in the open grassland decreased with elevation. Young growth originated from sexual reproduction at the lower forest but was comprised exclusively of root suckers at the treeline stand. We conclude that both the marked change in carbon allocation towards the root system, as well as the changes in root morphology with elevation indicate an adaptation to reduced nutrient supply under cold conditions of these Polylepis stands at the treeline in E Bolivia.

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Positive tree diversity effect on fine root biomass: via density dependence rather than spatial root partitioning

The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient-rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient-rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient-rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density-dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed.     

Environmental and Biotic Controls over Aboveground Biomass Throughout a Tropical Rain Forest

The environmental and biotic factors affecting spatial variation in canopy three-dimensional (3-D) structure and aboveground tree biomass (AGB) are poorly understood in tropical rain forests. We combined field measurements and airborne light detection and ranging (lidar) to quantify 3-D structure and AGB across a 5,016 ha rain forest reserve on the northeastern flank of Mauna Kea volcano, Hawaii Island. We compared AGB among native stands dominated by Metrosideros polymorpha found along a 600–1800 m elevation/climate gradient, and on three substrate-age classes of 5, 20, and 65 kyr. We also analyzed how alien tree invasion, canopy species dominance and topographic relief influence AGB levels. Canopy vertical profiles derived from lidar measurements were strong predictors (r ² = 0.78) of AGB across sites and species. Mean AGB ranged from 48 to 363 Mg ha⁻¹ in native forest stands. Increasing elevation corresponded to a 53–84% decrease in AGB levels, depending upon substrate age. Holding climate constant, changes in substrate age from 5 to 65 kyr corresponded to a 23–53% decline in biomass. Invasion by Psidium cattleianum and Ficus rubiginosa trees resulted in a 19–38% decrease in AGB, with these carbon losses mediated by substrate age. In contrast, the spread of former plantation tree species Fraxinus uhdei corresponded to a 7- to 10-fold increase in biomass. The effects of topographic relief at both local and regional scales were evident in the AGB maps, with poorly drained terrain harboring 76% lower biomass than forests on well-drained relief. Our results quantify the absolute and relative importance of environmental factors controlling spatial variation in tree biomass across a rain forest landscape, and highlight the rapid changes in carbon storage incurred following biological invasion. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  GPA and RFH conceived of or designed the study. GPA, RFH, TAV, DEK, and TKB performed research and analyzed data. GPA, RFH, DEK, and TKB contributed new methods or models. GPA wrote the article.     

地上竞争对林下红松生物量分配的影响

采用整株收获法研究林下红松地上、地下生物量分配特征及地上竞争对其生物量分配和生物量相对生长的影响。结果表明,(1)将整个树冠划分为等长的上、中、下三层,活枝生物量从上层到下层逐渐增加,而针叶生物量主要集中在树冠中下层且在中下层的分布无显著差异(P>0.05),随着地下生物量逐渐增加,小细根(<2 mm)、粗细根(2-5 mm)的比例逐渐减小,而粗根(>5 mm)所占比例逐渐增大;(2)地上竞争强度与胸径、树高呈显著指数相关(P<0.001),随着竞争强度增大,胸径和树高均逐渐减小,树高胸径比与竞争强度呈显著线性相关(P<0.05),而树冠比率与竞争强度之间无显著相关性(P>0.05);(3)随着竞争强度增大,树干生物量占整株生物量的相对比例逐渐减小,而细根(小细根和粗细根)生物量相对比例逐渐增大,活枝、针叶及粗根生物量相对比例与竞争强度相关性并不显著(P>0.05);(4)红松根冠比均值为0.15且根冠比并不受地上竞争的影响,茎叶比与竞争强度的相关性亦不显著(P>0.05);(5)地上竞争显著影响红松地上各器官生物量的相对生长,且竞争强度与生物量呈显著负相关(P<0.001)。