Multi-scale Drivers of Spatial Variation in Old-Growth Forest Carbon Density Disentangled with Lidar and an Individual-Based Landscape Model

Forest ecosystems are the most important terrestrial carbon (C) storage globally, and presently mitigate anthropogenic climate change by acting as a large and persistent sink for atmospheric CO₂. Yet, forest C density varies greatly in space, both globally and at stand and landscape levels. Understanding the multi-scale drivers of this variation is a prerequisite for robust and effective climate change mitigation in ecosystem management. Here, we used airborne light detection and ranging (Lidar) and a novel high-resolution simulation model of landscape dynamics (iLand) to identify the drivers of variation in C density for an old-growth forest landscape in Oregon, USA. With total ecosystem C in excess of 1?Gt?ha^?1 these ecosystems are among the most C-rich globally. Our findings revealed considerable spatial variability in stand-level C density across the landscape. Notwithstanding the distinct environmental gradients in our mountainous study area only 55.3% of this variation was explained by environmental drivers, with radiation and soil physical properties having a stronger influence than temperature and precipitation. The remaining variation in C stocks was largely attributable to emerging properties of stand dynamics (that is, stand structure and composition). Not only were density- and size-related indicators positively associated with C stocks but also diversity in composition and structure, documenting a close link between biodiversity and ecosystem functioning. We conclude that the complexity of old-growth forests contributes to their sustained high C levels, a finding that is relevant to managing forests for climate change mitigation.     

Dead Wood Necromass in a Moist Tropical Forest: Stocks,Fluxes, and Spatiotemporal Variability

Ecosystems - Woody debris (WD) stocks and fluxes are important components of forest carbon budgets and yet remain understudied, particularly in tropical forests. Here we present the most...     

Simulating Daily and Half-Hourly Fluxes of Forest Carbon Dioxide and Water Vapor Exchange with a Simple Model of Light and Water Use

For the large-scale application of simple, aggregated models, it is important to be able to link the values of model parameters to easily measurable ecosystem characteristics. However, the aggregation of model inputs and outputs over time and space can hamper this linkage. In this paper, two temporal versions of the same simple carbon dioxide (CO₂) and water exchange model, based on the concepts of water- and light-use efficiencies, were used to simulate the half-hourly and daily CO₂ and water exchange of a Douglas fir forest (Pseudotsuga menziesii (Mirb.) Franco) in the Netherlands for 2 years, before and after a thinning. We tested the performance of the models and the interpretability of changes in optimized parameter values, due to the thinning, in terms of ecosystem functioning. The performance of the half-hourly model was satisfactory, whereas the performance of the daily model was high for water exchange but clearly lower for CO₂ exchange. A comparison of the model parameters before and after the thinning showed that the coefficients of the half-hourly model could be separated into more physiologically determined and stand-determined characteristics, but this separation was not clear for the daily model. These results show that if the temporal resolution of the model is high enough, the effects of a major ecosystem manipulation, such as thinning, can be detected and interpreted using eddy flux data and a very simple biophysical model. The model parameters have an unambiguous interpretation and can be inferred from basic ecosystem observables, such as leaf area index (LAI) and aboveground biomass. A sensitivity analysis found strong correlations between parameter sets with similar model performance. For any comparison of the parameter values of different studies, ranges of parameter values and their correlations should be presented rather than one optimized value. Received 2 May 2001; accepted 15 February 2002.     

Leaf Water Repellency as an Adaptation to Tropical Montane Cloud Forest Environments

Adaptations that reduce water retention on leaf surfaces may increase photosynthetic capacity of cloud forests because carbon dioxide diffuses slower in water than air. Leaf water repellency was examined in three distinct ecosystems to test the hypothesis that tropical montane cloud forest species have a higher degree of leaf water repellency than species from tropical dry forests and species from temperate foothills-grassland vegetation. Leaf water repellency was measured by calculating the contact angle of the leaf surface and the line tangent to a water droplet through the point of contact on the adaxial and the abaxial surface. Leaf water repellency was significantly different between the three study areas. The hypothesis that leaf water repellency is higher in cloud forest species than tropical dry forests and temperate foothills-grassland vegetation was not confirmed in this study. Leaf water repellency was lower for cloud forest species (adaxial surface = 50.8°; abaxial surface = 82.9°) than tropical dry forest species (adaxial surface = 74.5°; abaxial surface = 87.3°) and temperate foothills-grassland species (adaxial surface = 77.6°; abaxial surface = 95.8°). The low values of leaf water repellency in cloud forest species may be influenced by presence of epiphylls and loss of epicuticular wax on the leaf surfaces.     

西双版纳热带山地雨林群落学特征分析

根据4块样地的调查资料,分析了西双版纳热带山地雨林的群落学特征。结果表明：在2500m^2的样地中,共计有维管束植物105～178种,分属于46～61科、80～130属。区系成分可分为11个类型,4块样地均以热带区系成分为主（约90％）,其中又以热带亚洲（印度-马来西亚）分布（23．40％～32．88％）和泛热带分布（25．20％～36．67％）的成分占优势。群落以高位芽植物为主,其中又以中（约50％）、小高位芽植物（21．30％～27．78％）占优势;在叶片特征上,群落以中叶（72．66％～82．54％）、单叶（75％～86．96％）、纸质（58．73％～62．22％）、全缘（76．67％～83．33％）、非尾尖（约90％）为主。低海拔带上山地雨林的热带区系成分以及藤本植物、中叶和全缘叶植物所占的比例高于高海拔带上的山地雨林,而小高位芽植物比例则低于后者,体现出较强的热带雨林性质。西双版纳的山地雨林与本地区的沟谷雨林、低丘雨林相比表现为热带区系成分、藤本植物、大高位芽植物以及巨叶、大叶植物较少,而温带成分、小高位芽植物、小叶植物较为丰富,体现出向常绿阔叶林的过渡特征。     

Aboveground vs. Belowground Carbon Stocks in African Tropical Lowland Rainforest: Drivers and Implications

BackgroundAfrican tropical rainforests are one of the most important hotspots to look for changes in the upcoming decades when it comes to C storage and release. The focus of studying C dynamics in these systems lies traditionally on living aboveground biomass. Belowground soil organic carbon stocks have received little attention and estimates of the size, controls and distribution of soil organic carbon stocks are highly uncertain. In our study on lowland rainforest in the central Congo basin, we combine both an assessment of the aboveground C stock with an assessment of the belowground C stock and analyze the latter in terms of functional pools and controlling factors.Conclusions/SignificanceWe suggest nutrient limitation, especially potassium, as the driver for aboveground versus belowground C allocation. However, other drivers such as C turnover, tree functional traits or demographic considerations cannot be excluded. We argue that large and unaccounted variability in C stocks is to be expected in African tropical rain-forests. Currently, these differences in aboveground and belowground C stocks are not adequately verified and implemented mechanistically into Earth System Models. This will, hence, introduce additional uncertainty to models and predictions of the response of C storage of the Congo basin forest to climate change and its contribution to the terrestrial C budget.     

Tree Species Composition Predicts Epiphytic Lichen Communities in an African Montane Rain Forest

The ecology of many tropical rain forest organisms, not the least in Africa, remains poorly understood. Here, we present a detailed ecological study of epiphytic lichens in the equatorial montane rain forest of Bwindi National Park (331 km²), Uganda. We evaluated all major lichen growth forms, including selected groups of crustose lichens. In 14 transects at elevations of 1290 m to 2500 m, we sampled 276 trees belonging to 60 species. We recorded all lichen species on each tree trunk between ground level and 2 m above the ground, yielding 191 lichen species in 67 genera, with a mean of 4.7 species per tree. We used non‐metric multi‐dimensional scaling to separate epiphytic lichen assemblages according to tree species composition and elevation. Structural equation modeling indicated that elevation influenced tree species composition and that tree species composition largely determined lichen species composition. Thus, elevation acted indirectly on the lichen assemblages. Further studies examining factors such as bark properties and lichen colonization ecology may clarify what determines the association between tree species and lichen assemblages. The link between lichen assemblages and large‐scale elevation patterns, as well as disturbance and regrowth histories, warrants further study. An analysis of lichen species composition on individual tree species that occur over large elevation ranges would distinguish the effect of tree species on lichen assemblages from the effect of elevation and thus climate. Our study highlights the limited extent of our knowledge of tropical epiphytic lichens.     

Photosynthesis-Nitrogen Relationships in Pioneer Plants of Disturbed Tropical Montane Forest Sites

Tropical forest disturbances lead to the establishment of secondary successional plant communities constituted by light demanding species with high relative growth rate that conduct to rapid canopy closure. Two main strategies for N nutrition are: (a) mineral N acquisition in the form of NH4 and NO3, and (b) symbiotic atmospheric N2 fixation. Given the high N requirement for maximization of leaf area and radiant energy absorption, we hypothesize that contrasting strategies of N nutrition in these environments are reflected in leaf photosynthetic characteristics. We compared the N-photosynthesis relationships and carbon balance parameters per unit leaf area as they vary with age in two species with contrasting N acquisition strategies: a N2-fixer Crotalaria anagyroides HBK (Papilionoideae), and a mineral-N user Verbesina turbacensis HBK (Asteraceae). N2 fixation capacity was associated to higher specific leaf area (SLA), higher photosynthetic capacity (Pmax) per unit leaf area and leaf mass, and higher N content per unit leaf mass. The N2-fixer species showed higher slope in the relationship Pmax-N per unit leaf mass and area when compared to the leaves of non-fixer species. Moreover, the intrinsic photosynthetic N use efficiency (Pmax/N) was higher in the N2 fixer than in leaves of the non-fixer species. Changes in N due to leaf age resulted in larger changes in CO2 flux density at the leaf level in the N2-fixer species. The higher photosynthetic capacity of the N2-fixer species was mechanistically related to higher stomatal conductance, internal CO2 concentration (ci) values closer to atmospheric CO2 concentration (ca), and lower intrinsic water use efficiency than the mineral N-user species. Despite their higher Pmax per unit leaf area, total non-structural saccharides concentration was lower in mature leaves of the N2-fixer plant as compared to the non-fixer counterpart. This might be caused by the presence of a larger root sink (symbionts) stimulating saccharides export and higher diurnal respiration rates.     

The Landscape Ecology of Tropical Secondary Forest in Montane Costa Rica

Multinomial logistic models of land use/land cover in montane Costa Rica and landscape pattern analysis showed that relative to agriculture, secondary forest occurred closer to old-growth forest, further from roads, in forest reserves, and at higher elevations. Collinearity between explanatory variables yielded simple multivariate models; proportion of surrounding old growth predicted secondary forest most accurately. An old-growth matrix [mean patch size (MPS) 24.5 ha], located mainly within protected areas, dominated elevations greater than 2500 m. A matrix of agriculture (MPS 23.5 ha), with smaller patches (approximately 9 ha) of secondary forest and old growth, dominated elevations from 1500 to 2500 m. Combining secondary forest with old growth decreased forest patch number and increased MPS from 7.3 to 37.1 ha. I concluded that: (a) secondary forest pattern is nonrandom, so ancillary data will aid its mapping with satellite imagery. The variables elevation, agriculture distance, road distance, and population density distinguished secondary forest from old growth with 74% accuracy; (b) socioeconomic and biological forces probably interact to create these secondary forest patterns; and (c) the strong association between secondary forest and old growth supports the concept that tropical forest recovery depends on the landscape structure of remnant forest. Received 16 February 1999; accepted 20 August 1999.     

Long-Term Impacts of Fuelwood Extraction on a Tropical Montane Cloud Forest

Fuelwood extracted from natural forests serves as a principal energy source in rural regions of many tropical countries. Although fuelwood extraction (even low intensities) might strongly impact the structure and species composition of natural forests, long-term studies remain scarce. Here, we estimate the potential long-term impacts (over several hundred years) of such repeated harvesting of single trees on tropical montane cloud forest in central Veracruz, Mexico, by applying a process-based forest growth model. We simulate a wide range of possible harvesting scenarios differing in wood volume harvested and preferred tree species and sizes, and use a set of indicators to compare their impacts on forest size structure and community composition. Results showed that the overall impact on forest structure and community composition increased linearly with the amount of harvested wood volume. Even at low levels of harvesting, forest size structure became more homogeneous in the long term because large old trees disappeared from the forest, but these changes might take decades or even centuries. Although recruitment of harvested species benefited from harvesting, species composition shifted to tree species that are not used for fuelwood. Our results demonstrate that fuelwood extraction can have marked long-term impacts on tropical montane cloud forests. The results also offer the possibility to support the design of management strategies for the natural species-rich forests that achieve a balance between economic needs and ecological goals of the stakeholders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Root Morphology and Anchorage of Six Native Tree Species from a Tropical Montane Forest and an Elfin Forest in Ecuador

Root architecture of tree species was investigated at two different altitudes in tropical forests in Ecuador. Increasing altitude was accompanied by higher wind speeds and more shallow soils, while slope angles of both sites were comparable (20–50°). Three tree species typical for the montane forest at 1900 m (Graffenrieda emarginata (Ruiz & Pav.) Triana (Melastomataceae), Clethra revoluta (Ruiz & Pav.) Spreng. (Clethraceae), Vismia tomentosa Ruiz & Pav. (Clusiaceae)) and for the elfin forest at 3000 m (Weinmannia loxensis Harling (Cunoniaceae), Clusia spec. (Clusiacaea) Styrax foveolaria Perkins (Styraceae)) were examined. At 1900 m, 92% of the trees grew upright, in comparison to 52% at 3000 m. At 3000 m, 48% of the trees were inclined, lying or even partly uprooted. At this altitude, all trees with tap roots or with shoots connected by coarse rhizomes, 83% of the trees with stilt roots, and 50% of the trees in which stems or roots were supported by other trees grew upright, suggesting that these characteristics were relevant for tree stability. Root system morphology differed markedly between altitudes. In contrast to 1900 m, where 20% of structural roots originated in the deeper mineral soil, root origin at 3000 m was restricted to the forest floor. The mean ratio of root cross sectional area to tree height decreased significantly from 6.1 × 10⁻³ m² m⁻¹ at 1900 m to 3.2 × 10⁻³ m² m⁻¹ at 3000 m. The extent of root asymmetry increased significantly from 0.29 at 1900 m to 0.62 at 3000 m. This was accompanied by a significantly lower number of dominant roots at 3000 m (2.3 compared to 3.8 at 1900 m). In conclusion, native tree species growing in tropical montane and elfin forests show a variety of root traits that improve tree stability. Root system asymmetry is less important for tree stability where anchorage is provided by a deep and solid root–soil plate. When deep rooting is impeded, root traits improving the horizontal extension of the root–soil plate are more pronounced or occur more frequently. Furthermore, mutual mechanical support of roots and stems of neighboring trees seems to be an appropriate mechanism to provide anchorage in soils with low bulk density and in environments with high wind speeds.     

Gross Primary Productivity of a High Elevation Tropical Montane Cloud Forest

For decades, the productivity of tropical montane cloud forests (TMCF) has been assumed to be lower than in tropical lowland forests due to nutrient limitation, lower temperatures, and frequent cloud immersion, although actual estimates of gross primary productivity (GPP) are very scarce. Here, we present the results of a process-based modeling estimate of GPP, using a soil–plant–atmosphere model, of a high elevation Peruvian TMCF. The model was parameterized with field-measured physiological and structural vegetation variables, and driven with meteorological data from the site. Modeled transpiration corroborated well with measured sap flow, and simulated GPP added up to 16.2 ± SE 1.6 Mg C ha^?1 y^?1. Dry season GPP was significantly lower than wet season GPP, although this difference was 17% and not caused by drought stress. The strongest environmental controls on simulated GPP were variation of photosynthetic active radiation and air temperature (T _air). Their relative importance likely varies with elevation and the local prevalence of cloud cover. Photosynthetic parameters (V _cmax and J _max) and leaf area index were the most important non-environmental controls on GPP. We additionally compared the modeled results with a recent estimate of GPP of the same Peruvian TMCF derived by the summing of ecosystem respiration and net productivity terms, which added up to 26 Mg C ha^?1 y^?1. Despite the uncertainties in modeling GPP we conclude that at this altitude GPP is, conservatively estimated, 30–40% lower than in lowland rainforest and this difference is driven mostly by cooler temperatures than changes in other parameters.     

大兴安岭天然落叶松林相容性立木含碳量模型研究

森林是生态系统的重要组成部分,是改善全球气候变暖趋势的机体,森林的固碳能力越来越受到重视,而研究森林中立木的含碳量对森林生态系统量化固碳能力具有重要意义。本文基于大兴安岭44株天然落叶松解析木实测数据和各器官碳密度样木数据,借鉴相容性生物量模型的思想来研究相容性立木含碳量模型,基于基础模型y=aD^b和y=a(D²H)^b,利用非线性度量误差模型系统构建了总含碳量和干、枝、叶、根四个分含碳量之间相容的一元和二元立木含碳量模型。通过比较各模型的拟合优度和独立检验统计量,计算拟合优度结果显示：在一元和二元基础模型下的4个器官,树干的确定系数R2分别为0.960,0.985,都是2个模型中各器官确定系数的最高值,树枝、树叶和树根相对偏低,但均达到了85%以上,说明建立的模型可行;模型检验统计量表明：一元和二元基础模型的树干模拟效率EF值为0.904,0.951,相应的预估精度P值为80.5%,85.5%,其次是树枝模拟效率0.830,0.898,精度都在70%以上,树叶和树根预估精度偏低,其值在70%左右。综上研究结果表明：二元立木含碳量预估模型的拟合及预测精度优于一元模型。     

Tropical Forest Fragmentation Affects Floral Visitors but Not the Structure of Individual-Based Palm-Pollinator Networks

Despite increasing knowledge about the effects of habitat loss on pollinators in natural landscapes, information is very limited regarding the underlying mechanisms of forest fragmentation affecting plant-pollinator interactions in such landscapes. Here, we used a network approach to describe the effects of forest fragmentation on the patterns of interactions involving the understory dominant palm Astrocaryum mexicanum (Arecaceae) and its floral visitors (including both effective and non-effective pollinators) at the individual level in a Mexican tropical rainforest landscape. Specifically, we asked: (i) Does fragment size affect the structure of individual-based plant-pollinator networks? (ii) Does the core of highly interacting visitor species change along the fragmentation size gradient? (iii) Does forest fragment size influence the abundance of effective pollinators of A. mexicanum? We found that fragment size did not affect the topological structure of the individual-based palm-pollinator network. Furthermore, while the composition of peripheral non-effective pollinators changed depending on fragment size, effective core generalist species of pollinators remained stable. We also observed that both abundance and variance of effective pollinators of male and female flowers of A. mexicanum increased with forest fragment size. These findings indicate that the presence of effective pollinators in the core of all forest fragments could keep the network structure stable along the gradient of forest fragmentation. In addition, pollination of A. mexicanum could be more effective in larger fragments, since the greater abundance of pollinators in these fragments may increase the amount of pollen and diversity of pollen donors between flowers of individual plants. Given the prevalence of fragmentation in tropical ecosystems, our results indicate that the current patterns of land use will have consequences on the underlying mechanisms of pollination in remnant forests.     

Carbon Stocks in an African Woodland Landscape: Spatial Distributions and Scales of Variation

Current knowledge of Africa’s carbon (C) pools is limited despite its importance in the global C budget. To increase the understanding of C stocks in African woodlands, we asked how C stocks in soil and vegetation vary across a miombo woodland landscape and to what degree and at what scales are these stocks linked? We sampled along a 5-km transect using a cyclic sampling scheme to allow geostatistical analyses. Soil C stocks in the top 5?cm (12.1?±?0.6?Mg?C?ha^?1 (±?SE)) and 30?cm depths (40.1?±?2.5?Mg?C?ha^?1) varied significantly at scales of a few meters (autocorrelation distance 14?m in 0–5-cm and 26?m in 0–30-cm interval), and aboveground (AG) woody C stocks (20.7?±?1.8?Mg?C?ha^?1) varied significantly at kilometer scales (1,426?m). Soil textural distributions were linked to topography (r ²?=?0.54) as were large-tree AG C stocks (r ²?=?0.70). AG C stocks were constrained to an upper boundary by soil texture with greater AG C being associated with coarser textured soils. Vegetation and soil C stocks were coupled in the landscape in the top 5?cm of soil (r ²?=?0.24) but not with deeper soil C stocks, which were coupled to soil clay content (r ²?=?0.38). This study is one of the most complete transect studies in an African miombo woodland, and suggests that C stock distributions are strongly linked to topography and soil texture. To optimize sampling strategies for C stock assessments in miombo, soil C should be sampled at more than 26?m apart, and AG C should be sampled at more than 1,426?m apart in plots larger than 0.5?ha.     

Forest Productivity and Efficiency of Resource Use Across a Chronosequence of Tropical Montane Soils

We tested the hypothesis that plants adjust to nutrient availability by altering carbon allocation patterns and nutrient-use efficiency (NUE = net primary production [NPP] per unit nutrient uptake), but are constrained by a trade-off between NUE and light-use efficiency () = NPP per unit intercepted light). NPP, NUE and ) were measured in montane Metrosideros polymorpha forest across a 4.1 x 10⁶ yr space for time substitution chronosequence in which available soil N and P pools change with site age. Although the range of N and P availability across sites was broad, there was little difference in NPP between sites, and in contrast to theories of carbon allocation relative to limiting resources, we found no consistent relationships in production allocation to leaves, fine roots or wood. However, canopy nutrient pools and fluxes were correlated with the mass of fine roots per unit soil volume and there was a weak but positive correlation of NPP with LAI. Patterns of ) and NUE across the soil developmental sequence were opposite to each other. ) increased as nutrient availability and nutrient turnover increased, while NUE decreased in response to the same influences but reached its highest values where either N or P availability and turnover of both N and P were low. A negative correlation between ) and NUE supports the hypothesis that a trade-off exists between ) and leaf characteristics affecting NUE.     

Structure of the Epiphyte Community in a Tropical Montane Forest in SW China

Vascular epiphytes are an understudied and particularly important component of tropical forest ecosystems. However, owing to the difficulties of access, little is known about the properties of epiphyte-host tree communities and the factors structuring them, especially in Asia. We investigated factors structuring the vascular epiphyte-host community and its network properties in a tropical montane forest in Xishuangbanna, SW China. Vascular epiphytes were surveyed in six plots located in mature forests. Six host and four micro-site environmental factors were investigated. Epiphyte diversity was strongly correlated with host size (DBH, diameter at breast height), while within hosts the highest epiphyte diversity was in the middle canopy and epiphyte diversity was significantly higher in sites with canopy soil or a moss mat than on bare bark. DBH, elevation and stem height explained 22% of the total variation in the epiphyte species assemblage among hosts, and DBH was the most important factor which alone explained 6% of the variation. Within hosts, 51% of the variation in epiphyte assemblage composition was explained by canopy position and substrate, and the most important single factor was substrate which accounted for 16% of the variation. Analysis of network properties indicated that the epiphyte host community was highly nested, with a low level of epiphyte specialization, and an almost even interaction strength between epiphytes and host trees. Together, these results indicate that large trees harbor a substantial proportion of the epiphyte community in this forest.     

Retention of Inorganic Nitrogen by Epiphytic Bryophytes in a Tropical Montane Forest1

We developed and evaluated a model of the canopy of a tropical montane forest at Monteverde, Costa Rica, to estimate inorganic nitrogen (N) retention by epiphytes from atmospheric deposition. We first estimated net retention of inorganic N by samples of epiphytic bryophytes, epiphyte assemblages, vascular epiphyte foliage, and host tree foliage that we exposed to cloud water and precipitation solutions. Results were then scaled up to the ecosystem level using a multilayered model of the canopy derived from measurements of forest structure and epiphyte mass. The model was driven with hourly meteorological and event‐based atmospheric deposition data, and model predictions were evaluated against measurements of throughfall collected at the site. Model predictions were similar to field measurements for both event‐based and annual hydrologic and inorganic N fluxes in throughfall. Simulation of individual events indicated that epiphytic bryophytes and epiphyte assemblages retained 33–67 percent of the inorganic N deposited in cloud water and precipitation. On an annual basis, the model predicted that epiphytic components retained 3.4 kg N ha/yr, equivalent to 50 percent of the inorganic N in atmospheric deposition (6.8 kg N ha/yr). Our results indicate that epiphytic bryophytes play a major role in N retention and cycling in this canopy by transforming highly mobile inorganic N (ca. 50% of atmospheric deposition is NO^?₃) to less mobile (exchangeable NH⁺₄) and recalcitrant forms in biomass and remaining litter and humus.