Immediate and long-term nitrogen oxide emissions from tropical forest soils exposed to elevated nitrogen input

Tropical nitrogen (N) deposition is projected to increase substantially within the coming decades. Increases in soil emissions of the climate‐relevant trace gases NO and N₂O are expected, but few studies address this possibility. We used N addition experiments to achieve N‐enriched conditions in contrasting montane and lowland forests and assessed changes in the timing and magnitude of soil N‐oxide emissions. We evaluated transitory effects, which occurred immediately after N addition, and long‐term effects measured at least 6 weeks after N addition. In the montane forest where stem growth was N limited, the first‐time N additions caused rapid increases in soil N‐oxide emissions. During the first 2 years of N addition, annual N‐oxide emissions were five times (transitory effect) and two times (long‐term effect) larger than controls. This contradicts the current assumption that N‐limited tropical montane forests will respond to N additions with only small and delayed increases in soil N‐oxide emissions. We attribute this fast and large response of soil N‐oxide emissions to the presence of an organic layer (a characteristic feature of this forest type) in which nitrification increased substantially following N addition. In the lowland forest where stem growth was neither N nor phosphorus (P) limited, the first‐time N additions caused only gradual and minimal increases in soil N‐oxide emissions. These first N additions were completed at the beginning of the wet season, and low soil water content may have limited nitrification. In contrast, the 9‐ and 10‐year N‐addition plots displayed instantaneous and large soil N‐oxide emissions. Annual N‐oxide emissions under chronic N addition were seven times (transitory effect) and four times (long‐term effect) larger than controls. Seasonal changes in soil water content also caused seasonal changes in soil N‐oxide emissions from the 9‐ and 10‐year N‐addition plots. This suggests that climate change scenarios, where rainfall quantity and seasonality change, will alter the relative importance of soil NO and N₂O emissions from tropical forests exposed to elevated N deposition.     

Nitrogen cycling in canopy soils of tropical montane forests responds rapidly to indirect N and P fertilization

Although the canopy can play an important role in forest nutrient cycles, canopy‐based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using ¹⁵N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH₄⁺ transformations decreased with increasing elevation; gross rates of NO₃^? transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient‐addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long‐term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.     

Latitudinal pattern of mountain vegetation zonation in southern and eastern Asia

A new scheme of altitudinal and latitudinal vegetation zonation is proposed for eastern Asia. The latitudinal patterns of mountain vegetation zonation show a clear boundary at ca. 20°–30° N. For the tropical mountains south of 20° N, the altitudinal series includes tropical lowland, tropical lower montane, and tropical upper montane zones. For the temperate mountains north of 30° N, the series includes temperate lowland, temperate lower montane, and temperate upper montane zones. The mountains located between 20° and 30° N show a transitional zonation pattern; the lower two zones are comparable to the lower two of the tropical zonation (tropical lowland and tropical lower montane), and the upper two zones are comparable to those of the temperate zonation (temperate lower montane and temperate upper montane). The tropical upper montane zone is not found north of 20°–30° N, while the tropical lower montane zone reaches down to sea level and constitutes the temperate lowland zone. Thus the zonation between 20° and 30° N includes tropical lowland, tropical lower montane/temperate lowland, temperate lower montane, and temperate upper montane zones. The latitudinal series of lowland rain forests follows the scheme of climatic division into tropical, subtropical/warm-temperate, cool-temperate and cold-temperate, with a shift of the respective life forms, evergreen, evergreen notophyllous, deciduous, and evergreen needle-leaved. The tropical lower montane forest can be correlated to the horizontal subtropical/ warm-temperate zone. The temperate altitudinal and latitudinal zonations above 30° N are correlated and show an inclined parallel pattern from high altitudes in the south to low altitudes down to sea level in the north.     

Substrate, climate, and land use controls over soil N dynamics and N-oxide emissions in Borneo

Nitrogen (N) enrichment of tropical ecosystems is likely to increase with rapid industrial and agricultural development, but the ecological consequences of N additions in these systems are not well understood. We measured soil N- oxide emissions and N transformations in primary rain forest ecosystems at four elevations and across two substrate types on Mt. Kinabalu, Borneo, before and after short-term experimental N additions. We also measured N pools and fluxes across a land use gradient of primary forest, burned secondary forest, and fertilized agriculture. Background soil N₂O and NO emissions in primary forest decreased with elevation, and soils derived from sedimentary substrates had larger pools of inorganic N, rates of nitrification, and N-oxide fluxes than ultrabasic soils when there were significant differences between substrate types. N-oxide emissions after N additions and background rates of nitrification were low in all soils derived from ultrabasic substrates compared to sedimentary substrates, even at lowland sites supporting, diverse Dipterocarp forests growing on morphologically similar Oxisols. Rates of potential nitrification were good predictors of N-oxide emissions after N additions. N₂O and NO fluxes were largest at low elevations and on sedimentary-derived soils compared to ultrabasic-derived soils, even at the smallest addition of N, 15kgNha^–1. Because current methods of soil classification do not explicitly characterize a number of soil chemical properties important to nutrient cycling, the use of soil maps to extrapolate biogeochemical processes to the region or globe may be limited in its accuracy and usefulness. In agricultural systems, management practices were more important than substrate type in controlling N-oxide emissions and soil N cycling. N-oxide fluxes from agricultural fields were more than an order of magnitude greater than from primary forests on the same substrate type and at the same elevation. As primary forests are cleared for intensive agriculture, soil N₂O and NO emissions are likely to far exceed those from the most N-saturated tropical forest ecosystems. This study highlights the inter-dependence of climate, substrate age, N deposition, and land-use practices determining N cycling and N-oxide emissions in humid tropical regions.     

Emissions of nitrous oxide from three tropical forests in Southern China in response to simulated nitrogen deposition

Emissions of nitrous oxide (N₂O) from the soil following simulated nitrogen (N) deposition in a disturbed (pine), a rehabilitated (pine and broadleaf mixed) and a mature (monsoon evergreen broadleaf) tropical forest in southern China were studied. The following hypotheses were tested: (1) addition of N will increase soil N₂O emission in tropical forests; and (2) any observed increase will be more pronounced in the mature forest than in the disturbed or rehabilitated forest due to the relatively high initial soil N concentration in the mature forest. The experiment was designed with four N treatment levels (three replicates; 0, 50, 100, 150 kg N ha⁻¹ year⁻¹ for C (Control), LN (Low-N), MN (Medium-N), and HN (High-N) treatment, respectively) in the mature forest, but only three levels in the disturbed and rehabilitated forests (C, LN and MN). Between October 2005 to September 2006, soil N₂O flux was measured using static chamber and gas chromatography methodology. Nitrogen had been applied previously to the plots since July 2003 and continued during soil N₂O flux measurement period. The annual mean rates of soil N₂O emission in the C plots were 24.1 ± 1.5, 26.2 ± 1.4, and 29.3 ± 1.6 μg N₂O–N m⁻² h⁻¹ in the disturbed, rehabilitated and mature forest, respectively. There was a significant increase in soil N₂O emission following N additions in the mature forest (38%, 41%, and 58% when compared to the C plots for the LN, MN, and HN plots, respectively). In the disturbed forest a significant increase (35%) was observed in the MN plots, but not in the LN plots. The rehabilitated forest showed no significant response to N additions. Increases in soil N₂O emission occurred primarily in the cool-dry season (November, December and January). Our results suggest that the response of soil N₂O emission to N deposition in tropical forests in southern China may vary depending on the soil N status and land-use history of the forest.     

模拟大气氮沉降对中国森林生态系统影响的研究进展

人类活动加剧了活性氮的生产和排放,并导致氮沉降日益增加并全球化。目前,人类活动对全球氮循环的干扰已经超出了地球系统安全运行的界限。中国已成为全球氮沉降的高发区域,高氮沉降已经威胁到生态系统的健康和安全,并成为生态文明建设过程中亟待理清和解决的热点问题。对国际上和中国森林生态系统模拟氮沉降研究的概况进行了综述,并从生物学和非生物学两大过程重点阐述模拟氮沉降增加对中国主要森林生态系统影响的研究进展。中国自2000年以后才开始重视大气氮沉降产生的生态环境问题,中国科学院华南植物园在国内森林生态系统模拟氮沉降试验研究上做出了开创性的贡献。模拟氮沉降研究表明,持续高氮输入将会显著改变森林生态系统的结构和功能,并威胁生态系统的健康发展,特别是处于氮沉降热点区域的中国中南部。森林生态系统的氮沉降效应依赖于系统的氮状态、土地利用历史、气候特征、林型和林龄等。最后,对未来的研究提出了一些建议,包括加强长期跟踪研究和不同气候带站点之间的联网研究,特别是在森林生态系统对长期氮沉降响应与适应的过程机制、地下碳氮吸存潜力研究、以及与其他全球变化因子的耦合研究等方面,以期为森林生态系统的可持续发展提供理论基础和管理依据。     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

Nitrogen deposition contributes to soil acidification in tropical ecosystems

Elevated anthropogenic nitrogen (N) deposition has greatly altered terrestrial ecosystem functioning, threatening ecosystem health via acidification and eutrophication in temperate and boreal forests across the northern hemisphere. However, response of forest soil acidification to N deposition has been less studied in humid tropics compared to other forest types. This study was designed to explore impacts of long‐term N deposition on soil acidification processes in tropical forests. We have established a long‐term N‐deposition experiment in an N‐rich lowland tropical forest of Southern China since 2002 with N addition as NH₄NO₃ of 0, 50, 100 and 150 kg N ha^?1 yr^?1. We measured soil acidification status and element leaching in soil drainage solution after 6‐year N addition. Results showed that our study site has been experiencing serious soil acidification and was quite acid‐sensitive showing high acidification (pH_(H2O)<4.0), negative water‐extracted acid neutralizing capacity (ANC) and low base saturation (BS,< 8%) throughout soil profiles. Long‐term N addition significantly accelerated soil acidification, leading to depleted base cations and decreased BS, and further lowered ANC. However, N addition did not alter exchangeable Al³⁺, but increased cation exchange capacity (CEC). Nitrogen addition‐induced increase in SOC is suggested to contribute to both higher CEC and lower pH. We further found that increased N addition greatly decreased soil solution pH at 20 cm depth, but not at 40 cm. Furthermore, there was no evidence that Al³⁺ was leaching out from the deeper soils. These unique responses in tropical climate likely resulted from: exchangeable H⁺ dominating changes of soil cation pool, an exhausted base cation pool, N‐addition stimulating SOC production, and N saturation. Our results suggest that long‐term N addition can contribute measurably to soil acidification, and that shortage of Ca and Mg should receive more attention than soil exchangeable Al in tropical forests with elevated N deposition in the future.     

Structural comparison of tropical montane rain forests along latitudinal and altitudinal gradients in south and east Asia

Geographical patterns of altitudinal zonation, floristic composition, and structural features of tropical montane rain forests were examined along latitudinal gradients in south and east Asia. On equatorial mountains, the tropical montane rain forests occur above 1000 m. Toward middle latitudes, they come farther down and reach sea level at c. 35° N. Thus, the forests are equivalent to the subtropical rain forests of the latitudinal, horizontal zonation series. They exhibit gradual changes in floristic composition and structure along both altitudinal and latitudinal gradients. On equatorial mountains, they are divided into three types, i.e. tropical lower montane, upper montane, and subalpine forests. The three tree regeneration types, having emergent, sporadic and inverse-J type stem-diameter class frequency distributions, coexist in the lower montane forests, but the upper and subalpine forests display only the inverse-J type species with a few species of the sporadic type. Toward the northern latitudinal limit, the distinction between the three tropical montane forest zones in equatorial mountains becomes less clear. This can be explained by temperature conditions: on equatorial mountains, a temperature sum of 85° C months which controls the upper limit of the lower montane forests, and a coldest month mean temperature of-1° C which controls the evergreen broad-leaved trees, appear at c. 2500 and c. 4000 m respectively. The altitudinal range between 2500 m and 3800 m, which is the upper forest limit, is covered by upper montane and subalpine forests. On the other hand, at the latitudinal northern limit, the tropical upper montane and subalpine forests cannot exist because the above mentioned two temperature conditions occur at nearly the same point. Thus, at the northern latitudinal limit of the tropical montane forests, the three zones of equatorial mountains amalgamate into a single subtropical lowland forest community. This is due to the seasonal temperature climate in middle latitudes in, e.g., central Japan and central China.A part of this paper was presented as an oral presentation at the Vth International Congress of Ecology, Yokohama 23–30.8.1990.     

In-depth analysis of N2O fluxes in tropical forest soils of the Congo Basin combining isotope and functional gene analysis

Primary tropical forests generally exhibit large gaseous nitrogen (N) losses, occurring as nitric oxide (NO), nitrous oxide (N₂O) or elemental nitrogen (N₂). The release of N₂O is of particular concern due to its high global warming potential and destruction of stratospheric ozone. Tropical forest soils are predicted to be among the largest natural sources of N₂O; however, despite being the world’s second-largest rainforest, measurements of gaseous N-losses from forest soils of the Congo Basin are scarce. In addition, long-term studies investigating N₂O fluxes from different forest ecosystem types (lowland and montane forests) are scarce. In this study we show that fluxes measured in the Congo Basin were lower than fluxes measured in the Neotropics, and in the tropical forests of Australia and South East Asia. In addition, we show that despite different climatic conditions, average annual N₂O fluxes in the Congo Basin’s lowland forests (0.97 ± 0.53 kg N ha⁻¹ year⁻¹) were comparable to those in its montane forest (0.88 ± 0.97 kg N ha⁻¹ year⁻¹). Measurements of soil pore air N₂O isotope data at multiple depths suggests that a microbial reduction of N₂O to N₂ within the soil may account for the observed low surface N₂O fluxes and low soil pore N₂O concentrations. The potential for microbial reduction is corroborated by a significant abundance and expression of the gene nosZ in soil samples from both study sites. Although isotopic and functional gene analyses indicate an enzymatic potential for complete denitrification, combined gaseous N-losses (N₂O, N₂) are unlikely to account for the missing N-sink in these forests. Other N-losses such as NO, N₂ via Feammox or hydrological particulate organic nitrogen export could play an important role in soils of the Congo Basin and should be the focus of future research.Subject terms: Microbiology, Biogeochemistry     

Response of nutrient dynamics of decomposing pine (Pinus massoniana) needles to simulated N deposition in a disturbed and a rehabilitated forest in tropical China

The effects of simulated N deposition on changes in mass, C, N and P of decomposing pine (Pinus massoniana) needles in a disturbed and a rehabilitated forest in tropical China were studied during a 24-month period. The objective of the study was to test the hypothesis that litter decomposition in a disturbed forest is more sensitive to N deposition rate than litter decomposition in a rehabilitated forest due to the relatively low nutrient status in the former as a result of constant human disturbance (harvesting understory and litter). The litterbag method and N treatments (control, no N addition; low-N, 5 g N m⁻² year⁻¹; medium-N, 10 g N m⁻² year⁻¹) were employed to evaluate decomposition. The results revealed that N addition increased (positive effect) mass loss rate and C release rate but suppressed (negative effect) the release rate of N and P from decomposing needles in both disturbed and rehabilitated forests. The enhanced needle decomposition rate by N addition was significantly related to the reduction in the C/N ratio in decomposing needles. However, N availability is not the sole factor limiting needle decomposition in both disturbed and rehabilitated forests. The positive effect was more sensitive to the N addition rate in the rehabilitated forest than in the disturbed forest, however the reverse was true for the negative effect. These results suggest that nutrient status could be one of the important factors in controlling the response of litter decomposition and its nutrient release to elevated N deposition in reforested ecosystems in the study region.     

海南霸王岭不同森林类型附生兰科植物的多样性和分布

附生兰科植物是热带林附生植物的主要类群之一,对于维持热带林生态系统的物种多样性及生态功能具有重要的作用。以海南岛霸王岭国家级自然保护区内的6种热带原始林类型(热带季雨林、低地雨林、热带针叶林、山地雨林、山地常绿林及山顶矮林)中的附生兰科植物为研究对象,通过样带调查(每个森林类型设置12个10m×50m的样带,记录每个样带内胸径(DBH)≥5cm的树木及藤本上附生兰科植物的物种名称、株数及附生位置)分析了附生兰科植物的物种多样性、附生位置及其在不同森林类型中的分布规律。结果表明:1)3.6hm2森林调查样带内共记录到附生兰科植物9634株,分属于26属60种;2)除趋势对应分析(DCA)结果表明,6种森林类型中的附生兰科植物可分成5组(其中,山地常绿林与山顶矮林内的附生兰科植物归为一组);3)分布海拔范围相临近的森林类型的附生兰科植物具有较高的相似性,山地常绿林和山顶矮林附生兰科植物的相似性最高(88.9%);4)6种森林类型中,较高海拔的3种森林类型(山地雨林、山地常绿林和山顶矮林)中,附生兰科植物的丰富度和多度均显著高于其在较低海拔的3种森林类型(热带季雨林、低地雨林和热带针叶林),其中,附生兰科植物在山地常绿林内的丰富度和多度均最高;5)热带季雨林、低地雨林、热带针叶林及山地雨林内,宿主冠区附生兰科植物的多度均高于干区;山地常绿林内两者之间无显著差异;而山顶矮林干区的附生兰科植物的多度高于冠区;6)调查木上附生兰科植物的发生率在高海拔森林类型均高于其在低海拔森林类型,各森林类型内附生兰科植物的多度及物种丰富度与宿主胸径均存在显著正相关关系。     

Throughfall exclusion and leaf litter addition drive higher rates of soil nitrous oxide emissions from a lowland wet tropical forest

Tropical forests are a significant global source of the greenhouse gas nitrous oxide (N₂O). Predicted environmental changes for this biome highlight the need to understand how simultaneous changes in precipitation and labile carbon (C) availability may affect soil N₂O production. We conducted a small‐scale throughfall and leaf litter manipulation in a lowland tropical forest in southwestern Costa Rica to test how potential changes in both water and litter derived labile C inputs to soils may alter N₂O emissions. Experimentally reducing throughfall in this wet tropical forest significantly increased soil emissions of N₂O, and our data suggest that at least part of this response was driven by an increase in the concentration of dissolved organic carbon [DOC] inputs delivered from litter to soil under the drier conditions. Furthermore, [DOC] was significantly correlated with N₂O emissions across both throughfall and litterfall manipulation plots, despite the fact that native NO₃^? pools in this site were generally small. Our results highlight the importance of understanding not only the potential direct effects of changing precipitation on soil biogeochemistry, but also the indirect effects resulting from interactions between the hydrologic, C and N cycles. Finally, over all sampling events we observed lower mean N₂O emissions (<1 ng N₂O‐N cm^?2 h^?1) than reported for many other lowland tropical forests, perhaps reflecting a more general pattern of increasing relative N constraints to biological activity as one moves from drier to wetter portions of the lowland tropical forest biome.     

Canopy soil greenhouse gas dynamics in response to indirect fertilization across an elevation gradient of tropical montane forests

Canopy soils can significantly contribute to aboveground labile biomass, especially in tropical montane forests. Whether they also contribute to the exchange of greenhouse gases is unknown. To examine the importance of canopy soils to tropical forest‐soil greenhouse gas exchange, we quantified gas fluxes from canopy soil cores along an elevation gradient with 4 yr of nutrient addition to the forest floor. Canopy soil contributed 5–12 percent of combined (canopy + forest floor) soil CO₂ emissions but CH₄ and N₂O fluxes were low. At 2000 m, phosphorus decreased CO₂ emissions (>40%) and nitrogen slightly increased CH₄ uptake and N₂O emissions. Our results show that canopy soils may contribute significantly to combined soil greenhouse gas fluxes in montane regions with high accumulations of canopy soil. We also show that changes in fluxes could occur with chronic nutrient deposition.     

Are Montane Forests Demographic Sinks for Bornean White‐bearded Gibbons Hylobates albibarbis?

I estimated habitat-specific population densities for a population of Bornean white-bearded gibbons Hylobates albibarbis inhabiting seven distinct forest types at Gunung Palung National Park, West Kalimantan, Indonesia. Population densities in montane forests (0.44 individuals/km²) were almost ten times lower than those in the next best habitat (upland granite forest; 4.2 individuals/km²) and far lower than those in lowland forest types. Demographic data on 33 gibbon groups living across the seven forest types showed that reproduction was substantially depressed in montane forests compared to high-quality lowland habitats. A simple model suggests that montane forests are demographic sinks for gibbons at Gunung Palung. Follow-up data from observations of montane groups 5 yr after the initial observation period support this result. As high-quality lowland forests (source habitat for gibbons) are being disproportionately lost in and around Gunung Palung National Park due to illegal logging and conversion to oil palm plantations, an increasing percentage of the remaining forest in the park comprises sink habitat for gibbons. This result has disquieting implications for the long-term viability of gibbon populations at Gunung Palung. In addition, as montane forests are generally low-quality habitat for most rainforest vertebrates, and since lowland forests are being lost at alarming rates across the tropics, source-sink population dynamics similar to those I describe here may characterize populations of other tropical vertebrate species.     

Extraradical hyphae exhibit more plastic nutrient-acquisition strategies than roots under nitrogen enrichment in ectomycorrhiza-dominated forests

Ectomycorrhizal (ECM) functional traits related to nutrient acquisition are impacted by nitrogen (N) deposition. However, less is known about whether these nutrient-acquisition traits associated with roots and hyphae differentially respond to increased N deposition in ECM-dominated forests with different initial N status. We conducted a chronic N addition experiment (25 kg N ha⁻¹ year⁻¹) in two ECM-dominated forests with contrasting initial N status, that is, a Pinus armandii forest (with relatively low N availability) and a Picea asperata forest (with relatively high N availability), to assess nutrient-mining and nutrient-foraging strategies associated with roots and hyphae under N addition. We show that nutrient-acquisition strategies of roots and hyphae differently respond to increased N addition. Root nutrient-acquisition strategies showed a consistent response to N addition, regardless of initial forest nutrient status, shifting from organic N mining toward inorganic N foraging. In contrast, the hyphal nutrient-acquisition strategy showed diverse responses to N addition depending on initial forest N status. In the Pinus armandii forest, trees increased belowground carbon (C) allocation to ECM fungi thus enhancing hyphal N-mining capacity under increased N availability. By comparison, in the Picea asperata forest, ECM fungi enhanced both capacities of P foraging and P mining in response to N-induced P limitation. In conclusion, our results demonstrate that ECM fungal hyphae exhibit greater plasticity in nutrient-mining and nutrient-foraging strategies than roots do in response to changes of nutrient status induced by N deposition. This study highlights the importance of ECM associations in tree acclimation and forest function stability under changing environments.     

On the Classification of Forest Vegetation in Xishuangbanna,Southern Yunnnan

Xishuangbanna of southern Yunnan is a region of extremely interest to biologists and also a hotspot for biodiversity conservation . It is located in a transitional zone from tropical Southeast Asia to temperate East Asia biogeographically. The present paper reviewed vegetation types of Xishuangbanna and suggested a revised classification system based on theupdated study results over the last two decades . By combining physiognomic and floristic characteristics with ecological performances and habitats , the primary forest vegetation in Xishuangbanna can be organized into four main vegetation types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad-leaved forest and tropical monsoon forest. The tropical rain forest can be classified into two subtypes , i. e. tropical seasonal rain forest in the lowlands and tropical montane rain forest on higher elevations. The tropical seasonal rain forest in this region shows similar forest profile and physiognomic characteristics to those of equatorial lowland rain forests and is a type of world tropical rain forest. Because of conspicuous similarity on floristic composition , the tropical seasonal rain forest in Xishuangbanna is a type of tropical Asian rain forest . However , since the tropical seasonal rain forest occurs at the northern edge of tropical SE Asia, it differs from typical lowland rain forests in equatorial areas in maintaining some deciduous trees in the canopy layer , fewer megaphanerophytes and epiphytes but more abundant lianas and more plants with microphyll . It is a type of semi-evergreen rain forest at the northern edge of the tropical zone . The tropical montane rain forest occurs in wet montane habitats and is similar to the lower montane rain forests in equatorial Asia in floristic composition and physiognomy . It is a variety of lower montane rain forests at the northern tropical edges of tropical rain forests . The tropical seasonal moist forest occurs on middle and upper limestone mountains and is similar to the tropical montane evergreen broad-leaved forest of the region in physiognomy, but it differs from the latter in floristic composition. The monsoon forest in Xishuangbanna is a tropical deciduous forest under the influence of a strong monsoon climate and is considered to be a transitional vegetation type between tropical rain forest and savanna in physiognomy and distribution. The tropical montane evergreen broad- leaved forest is the main vegetation type in mountain areas . It is dominated by the tree species of Fagaceae , Euphorbiaceae , Theaceae and Lauraceae in majority. It differs from the tropical montane rain forests in lack of epiphytes and having more abundant lianas and plants with compound leaves . It is considered to be a distinct vegetation type in the northern margin of mainland southeastern Asia controlling by a strong monsoon climate, based on its floristic and physiognomic characteristics.     

论滇南西双版纳的森林植被分类

本文基于多年研究成果的总结,对西双版纳森林植被的分类、主要植被类型及其特征进行了系统归纳,并讨论了它们与世界类似热带森林植被的关系。以群落的生态外貌与结构、种类组成和生境特征相结合作为植被分类的原则和依据,可以将西双版纳的热带森林植被分类为热带雨林、热带季节性湿润林、热带季雨林和热带山地常绿阔叶林四个主要的植被型,包括有至少二十个群系。热带雨林包括热带季节雨林和热带山地(低山)雨林二个植被亚型。热带季节雨林具有与赤道低地热带雨林几乎一样的群落结构和生态外貌特征,是亚洲热带雨林的一个类型,但由于发生在季风热带北缘纬度和海拔的极限条件下,受到季节性干旱和热量不足的影响,在其林冠层中有一定比例的落叶树种存在,大高位芽植物和附生植物较逊色而藤本植物和在叶级谱上的小叶型植物更丰富,这些特征又有别于赤道低地的热带雨林。热带山地雨林是热带雨林的山地亚型,是该地区热带山地较湿润生境的一种森林类型,它在植物区系组成和生态外貌特征上类似于热带亚洲的低山雨林,隶属于广义热带雨林植被型下的低山雨林亚型。热带季节性湿润林分布在石灰岩山坡中、上部,在群落外貌上类似热带山地常绿阔叶林但在植物区系组成上与后者不同,它是石灰岩山地垂直带上的一种植被类型。热带季雨林是分布在该地区开阔河谷盆地及河岸受季风影响强烈的生境的一种热带落叶森林,是介于热带雨林与萨王纳之间的植被类型。热带山地常绿阔叶林(季风常绿阔叶林)是西双版纳的主要山地植被类型,它分布在热带季节雨林带之上偏干的山地生境。它在植物区系组成上不同于该地区的热带季节雨林,在生态外貌特征上亦不同于热带山地雨林,是发育在受地区性季风气候强烈影响的热带山地的一种森林植被类型。     

Carbon and nutrient limitation of soil microorganisms and microbial grazers in a tropical montane rain forest

We investigated the role of carbon, nitrogen and phosphorus as limiting factors of microorganisms and microbial grazers (testate amoebae) in a montane tropical rain forest in southern Ecuador. Carbon (as glucose), nitrogen (as NH₄NO₃) and phosphorus (as NaH₂PO₄) were added separately and in combination bimonthly to experimental plots for 20 months. By adding glucose and nutrients we expected to increase the growth of microorganisms as the major food resource of testate amoebae. The response of microorganisms to experimental treatments was determined by analysing microbial biomass (SIR), fungal biomass and microbial community composition as measured by phospholipid fatty acids (PLFAs). We hypothesized that the response of testate amoebae is closely linked to that of microorganisms. Carbon addition strongly increased ergosterol concentration and, less pronounced, the amount of linoleic acid as fungal biomarker, suggesting that saprotrophic fungi are limited by carbon. Microbial biomass and ergosterol concentrations reached a maximum in the combined treatment with C, N and P indicating that both N and P also were in short supply. In contrast to saprotrophic fungi and microorganisms in total, testate amoebae suffered from the addition of C and reached maximum density by the addition of N. The results indicate that saprotrophic fungi in tropical montane rain forests are mainly limited by carbon whereas gram positive and negative bacteria benefit from increased availability of P. Testate amoebae suffered from increased dominance of saprotrophic fungi in glucose treatments but benefited from increased supply of N. The results show that testate amoebae of tropical montane rain forests are controlled by bottom–up forces relying on specific food resources rather than the amount of bacterial biomass with saprotrophic fungi functioning as major antagonists. Compared to temperate systems microbial food webs in tropical forests therefore may be much more complex than previously assumed with trophic links being rather specific and antagonistic interactions overriding trophic interactions.     

Relationships among precipitation regime, nutrient availability, and carbon turnover in tropical rain forests

The effect of high precipitation regime in tropical forests is poorly known despite indications of its potentially negative effects on nutrient availability and carbon (C) cycling. Our goal was to determine if there was an effect of high rainfall on nitrogen (N) and phosphorous (P) availability and indexes of C cycling in lowland tropical rain forests exposed to a broad range of mean annual precipitation (MAP). We predicted that C turnover time would increase with MAP while the availability of N and P would decrease. We studied seven Neotropical lowland forests covering a MAP range between 2,700 and 9,500 mm. We used radiocarbon (?¹⁴C) from the atmosphere and respired from soil organic matter to estimate residence time of C in plants and soils. We also used C, N, and P concentrations and the stable isotope ratio of N (δ¹⁵N) in live and dead plant tissues and in soils as proxies for nutrient availability. Negative δ¹⁵N values indicated that the wettest forests had N cycles that did not exhibit isotope-fractionating losses and were potentially N-limited. Element ratios (N:P and C:P) in senescent leaves, litter, and live roots showed that P resorption increased considerably with MAP, which points towards increasing P-limitation under high MAP regimes. Soil C content increased with MAP but C turnover time only showed a weak relationship with MAP, probably due to variations in soil parent material and age along the MAP gradient. In contrast, comparing C turnover directly to nutrient availability showed strong relationships between C turnover time, N availability (δ¹⁵N), and P availability (N:P) in senescent leaves and litter. Thus, an effect of MAP on carbon cycling appeared to be indirectly mediated by nutrient availability. Our results suggest that soil nutrient availability plays a central role in the dynamic of C cycling in tropical rain forests.