Leaf-litter decomposition of 15 tree species in a lowland tropical rain forest in Sarawak: decomposition rates and initial litter chemistry

In a lowland tropical rain forest in Sarawak, leaf-litter decomposition and the initial litter chemistry of 15 tree species were studied. During 13 months of field experiment, weight loss of litter samples was between 44% and 91%, and calculated decomposition rate constants (k) ranged from 0.38 to 2.36 year⁻¹. The initial litter chemistry also varied widely (coefficients of variation: 19%–74%) and showed low N and P concentrations and high acid-insoluble residue (AIS) concentration. For nutrient-related litter chemistry, correlations with the decomposition rate were significant only for P concentration, C/P ratio, and AIS/P ratio (r _s = 0.59, −0.62, and −0.68, n = 15, P < 0.05, respectively). For organic constituents, correlations were significant for concentrations of AIS and total carbohydrates, and AIS/acid-soluble carbohydrate ratio (r _s = −0.81, 0.51, and −0.76, n = 15, P < 0.05, respectively). These results suggested that the relatively slow mean rate of decomposition (k = 1.10) was presumably due to the low litter quality (low P concentration and high AIS concentration), and that P might influence the decomposition rate; but organic constituents, especially the concentration of AIS, were more important components of initial litter chemistry than nutrient concentrations.     

Leaf litter decomposition of dominant tree species of Namdapha National Park,Arunachal Pradesh,northeast India

Rates of weight loss and nutrient (N and P) release patterns were studied in the leaf litter of the dominant tree species (Ailanthus grandis, Altingia excelsa, Castanopsis indica, Duabanga sonneriatioides, Dysoxylum binectariferum, Mesua ferrea, Shorea assamica, Taluma hodgsonii, Terminalia myriocarpa and Vatica lancefolia) of a tropical wet evergreen forest of northeast India. Nitrogen and phosphorus mineralization rate and decay pattern varied significantly from species to species. In general, the decay pattern, characterized by using a composite polynomial regression equation, exhibited three distinct phases of decay during litter decomposition—an initial slow decay phase (0.063% weight loss day⁻¹), followed by a rapid decay phase (0.494% weight loss day⁻¹) and a final slow decay phase (0.136% weight loss day⁻¹). The initial chemical composition of the litter affected decomposition rates and patterns. Species like D. sonneriatoides, D. binectariferum, and T. hodgsonii with higher N and P content, lower carbon and lignin content, and lower C:N ratio and lignin:N ratio exhibited relatively faster decomposition rates than the other species, for example M. ferrea, C. indica and A. grandis. A slow decay rate was recorded for species such as M. ferrea, C. indica, and A. grandis. The initial N and P content of litter showed significant positive correlations with decay rates. Carbon and lignin content, lignin:N, and C:N showed significant negative correlations with decay rates. Soil total N and P, and rainfall, soil temperature, and soil moisture had positive correlations with decay rates. The rapid decomposition rates observed in comparison with other different forest litter decay rates confirm that tropical wet evergreen forest species are characterized by faster decomposition rates, indicating a faster rate of organic matter turnover and rapid nutrient cycling.     

Contribution of agroforestry trees to nutrient requirements of intercropped plants

A major tenet of agroforestry, that trees maintain soil fertility, is based primarily on observations of higher crop yields near trees or where trees were previously grown. Recently objective analyses and controlled experiments have addressed this topic. This paper examines the issues of tree prunings containing sufficient nutrients to meet crop demands, the timing of nutrient transfer from decomposition to intercrops, the percent of nutrients released that are taken up by the crop, and the fate of nutrients not taken up by the crop.The amount of nutrients provided by prunings are determined by the production rate and nutrient concentrations, both depending on climate, soil type, tree species, plant part, tree density and tree pruning regime. A large number of screening and alley cropping trials in different climate-soil environments indicate that prunings of several tree species contain sufficient nutrients to meet crop demand, with the notable exception of phosphorus. Specific recommendations for the appropriate trees in a given environment await synthesis of existing data, currently only general guidelines can be provided.Tree biomass containing sufficient nutrients to meet crop demand is not enough, the nutrients must be supplied in synchrony to crop needs. Nutrient release patterns from organic materials are, in part, determined by their chemical composition, or quality. Leguminous materials release nitrogen immediately, unless they contain high levels of lignin or polyphenols. Nonlegumes and litter of both legumes and nonlegumes generally immobilize N initially. There is little data on release patterns of other nutrients. Indices that predict nutrient release patterns will assist in the selection of species for synchronizing with crop demand and improve nutrient use-efficiency.Field trials with agroforestry species ranging in quality show that as much as 80% of the nutrients are released during the course of annual crop growth but less than 20% is captured by the crop, a low nutrient-use efficiency. There are insufficient data to determine how much of the N not captured by the crop is captured by the trees or is in the soil organic matter, the availability of that N to subsequent crops, or how much of that N is lost through leaching, volatilization or denitrification. Longer term trials are needed.     

Litterfall, litter decomposition and nutrient dynamics in a subtropical natural oak forest and managed plantation in northeastern India

In northeastern India, subtropical forests are over-exploited for timber, fuel wood and common agricultural practice like shifting cultivation, which are responsible for the degradation of natural forest. In degraded areas, large-scale plantations of different species of Quercus have been raised since 1980 for the production of economic Tasar silk. Conversion of natural forest into plantation affects the process of nutrient cycling due to management practices. Thus, it would be of importance to study the litterfall, litter decomposition process and the factors regulating the rate of litter decay in these ecosystems to improve recommendations for their management and conservation. We recorded litterfall by using litter traps and decomposition of leaf litter by nylon net bag technique to understand the amount of organic matter and nutrient return and their release in soils of forest and plantation in Manipur, northeast India. Total litterfall was 419.9 g m⁻² year⁻¹ in plantation and 547.7 g m⁻² year⁻¹ in forest. Litter decomposition rate was faster at plantation site than the forest in the early stage of litter decomposition whereas the reverse was observed at later stages of decomposition. Stepwise regression analysis showed the significant role of relative humidity and mean temperature on mass loss rates in the forest. Relative humidity, maximum temperature, population of fungi and actinomycetes were the best predictor variables for mass loss rates in plantation. Nutrient retranslocation efficiency and the immobilization of N and P in forest litter were higher than plantation. This suggests that Q. serrata growing in natural ecosystem in oligotrophic condition adapted strong nutrient conservation mechanisms to compete with the other plant species for the meager soil nutrients. The same species in plantation loses these adaptive capabilities because of exogenous supply of nutrients and in the absence of intense competition with other plant species. Thus, the optimization of organic and chemical fertilizer input in plantation is recommended for maintaining the soil fertility level to produce quality leaf for silkworm by conserving essential nutrients in the system.     

滇南3种不同类型热带森林优势种幼树树干生态化学计量特征

目的 探讨滇南地区热带、亚热带森林物种养分元素分配模式、受限制状况、吸收利用资源能力等特征,以期揭示外界环境变化下物种的适应性机制。 方法 以滇南区域热带季节雨林、热带山地雨林和中山湿性常绿阔叶林3种热带森林主要优势种为研究对象,分析不同类型森林不同层次优势种幼树树干C、N、P化学计量特征变化。 结果 1）3种森林优势种幼树树干全C含量随海拔梯度的增加总体呈增加趋势,全N含量和全P含量随海拔梯度的增加呈降低趋势;2）中山湿性常绿阔叶林的C:N、C:P相对较高,热带季节雨林和热带山地雨林N:P均小于14,中山湿性常绿阔叶林大于16;3）森林上层优势种幼树树干全C、全N含量均低于下层树种,全P含量则高于下层树种,上层优势种N:P均小于14,下层优势种大于16,说明上层优势种生长主要受到N元素的限制,且吸收和利用养分能力高于下层优势种;4）冗余分析表明：海拔和层次对优势种幼树树干养分含量及化学计量比的影响较大。 结论 滇南地区3种森林中,中山湿性常绿阔叶林拥有相对较高的氮、磷利用效率,本区生态系统主要受到N、P元素的共同限制作用。不同生长环境条件下植物体内养分元素生态化学计量特征的差异性,可以进一步揭示物种适应各种生态环境的应对机制。     

Phosphorus and base cation accumulation and release patterns in decomposing Scots pine,Norway spruce and silver birch stumps

Tree stumps are integral constituents of managed forest ecosystems, but their role in nutrient cycling is poorly understood. We studied phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) dynamics in decomposing Scots pine (Pinus sylvestris), Norway spruce (Picea abies), and silver birch (Betula pendula) stumps in southern Finland in a chronosequence of 0-, 5-, 10-, 20-, 30- and 40-year-old clear-cut areas. Along with the decomposition of pine and spruce stumps, the amount of P in stumps increased, but K and Ca were released, and the amount of Mg initially decreased and then increased. All nutrients, except K, accumulated in birch stumps during the first ten years, but were released thereafter. After 40 years of decomposition, pine and spruce stumps contained 180% and 202% of their initial amounts of P, respectively. In addition, the amounts of Mg were larger than the initial amounts in 40-year decomposed pine (126%) and spruce (202%) stumps. In contrast, birch stumps lost 64% and 75%, respectively, of their initial amounts of P and Mg over a 40-year period. The stumps of all the species were found to release K and Ca. Pine, spruce and birch stumps lost 48%, 64% and 87% of their initial amount of K, and 49%, 35% and 42% of their initial amount of Ca, respectively, during the 40-year period. The results indicate that decomposing stumps of the major tree species in Fennoscandian forests are long-term nutrient pools and they serve as P sinks, thus potentially reducing P leaching after clear-cutting.     

Distribution of Selected Timber Species of a Central African Rain Forest in Relation to Topography and Soil Heterogeneity: Implications for Forest Management

Several studies have looked at the relative importance of habitat association of tree species in maintaining tropical tree species diversity at a variety of scales including the importance of edaphic factors. The objective of this study was to test hypotheses on the distribution of important timber species in relation to topographic position and soil chemical parameters. Tree species from a Central African rain forest were selected within a 100-ha plot in southwestern Central Africa Republic. Individuals of Lovoa trichilioides, Nesogordonia papaverifera, Pterocarpus soyauxii, Terminalia superba ≥ 10 cm diameter at breast height (dbh) were inventoried then mapped to topography using a survey laser. A torus translation was performed to test for associations between tree species distribution and topographic position as well as soil mineral nutrients for three of the four species. Trees ≥ 30 cm dbh of all three species showed no association with topographic position. However, the distribution of T. superba was significantly associated with high concentrations of soil mineral nutrients. Evidence from mineral nutrient levels indicates that extrapolating topography to define habitat and soil type can be very misleading. Because many of Central Africa's most important timber species have distributions linked to edaphic factors, it is clear that soils must be considered equally as much as topographic position for management plans of logging concessions within Central African tropical forests.     

海南岛霸王岭热带山地雨林林隙更新生态位的研究

通过对海南岛霸王岭热带山地雨林林隙更新生态位的研究表明：同一树种在不同大小和年龄级林隙内的优势度不同，不同树种在同一大小和年龄级林隙内的优势度不同。根据树种在不同大小级林隙内的生态位宽度的变化，可分为生态位幅度宽、中等、较小和狭小4类；根据树种在不同年龄级林隙内的生态位宽度的变化，可分为对林隙时间生态资源利用充分、利用较充分、利用不充分和利用很少的4类树种。不同的树种或树种组对不同大小和年龄阶段林隙生态资源利用的不一致性，表明热带山地雨林中不同树种的林隙大小级生态位和林隙时间段生态位是相对分离的。分析表明，海南岛热带山地雨林中有很大一部分树种是属于低密度、狭生态位的树种。从物种多样性的角度来看，大量低密度、狭生态位树种的存在表明热带山地雨林是一种较为脆弱的生态系统类型。     

Decomposition and nutrient release in needle litter from nitrogen‐fertilized scots pine (pinus sylvestris) stands

Decomposition of Scots pine needle litter originating from five stands treated with different amounts of nitrogen fertilizer was measured over a 4‐year period in a mature Scots pine forest. The litter types, which differed in initial concentrations of nitrogen, phosphorus, potassium, and sulfur, but not in gross organic composition, were studied with respect to mass loss, ingrowth of total fungal mycelium, and net release of nutrients. During the first year of decomposition, rates of mass loss and ingrowth of fungal mycelium were highest in the nutrient‐rich litter. Phosphorus concentration was found to be the main factor affecting mass‐loss rate, and the rate of fungal ingrowth was positively correlated with initial nitrogen concentration. After this initial period, decomposition rates decreased, and after 4 years, accumulated mass loss and amounts of fungal mycelium were similar in all five litter types. These findings may be due, in part, to a lower rate of lignin decomposition in nitrogen‐enriched litter. Of the elements, potassium and magnesium were most rapidly lost from the litter, and their release was most pronounced during the first year. Calcium release was proportional to the loss in organic matter. Initially, the release of nitrogen and phosphorus was positively related to their concentrations in litter, however, during later stages of decomposition the differences among litter types levelled out. There was a tendency for concentrations of all elements, except nitrogen, in the different litters to approach similar levels as decomposition proceeded. Thus, after 4 years the nutrient composition of the various litter types was very similar, except for higher nitrogen concentrations in the originally most nutrient‐rich litters. The importance of the results in terms of substrate quality in fertilized as well as in non‐fertilized forests is discussed.     

Impacts of mixed litter decomposition from Robinia pseudoacacia and other tree species on C loss and nutrient release in the Loess Plateau of China

The productivity of Robinia pseudoacacia(R.p.) pure forest usually declines at the late growth stage,and reforming it into mixed forests could be a promising way to resolve this problem. When choosing a suitable tree species that can be mixed with R.p., the interspecific relationship is an important issue. Therefore, we gathered the autumn litter fall from R.p. and 10 other species from the Loess Plateau of China were mixed in dual species litterbags(R.p.+each other species) and buried them in soil for a 345 days lab decay incubation. We measured the litter mass loss and nutrient contents to determine whether the nutrient release was affected by mixed species litter decomposition. The impacts of mixed litter decomposition on macro-elements release were more obvious than on micro-elements. The litters with similar substrate quality might show variable impacts on nutrients release in mixed decomposition. The C loss and release of nutrient was improved by descending order when R.p. litter was mixed with Hippophae rhamnoides, Ulmus pumila, Populus simonii, Larix principis-rupprechtii and Quercus liaotungensis(Q.l.). But, except for Q.l., only the other species were recommended as suitable mix-plants for R.p. since promoting a high turnover of the nutrient in the litter compartment and a rapid availability for tree.     

Decomposition and nutrient release of four potential mulching materials for poplar plantations on upland sites

The application of organic residues and the release dynamics of nutrients during decomposition may be crucial to improving poplar growth in upland areas. Field experiments were conducted in Guizhou Province, southwestern China, to determine differences in biomass decomposition of four potential mulching materials in one-year-old poplar plantations. Results showed the initial total content of N, P, K, Ca and Mg in the mulches ranged from 30.0 to 35.0 g/kg and was significantly greater in Quercus fabri, Imperata cylindrica var. major and Pteridium aquilinum var. latiuscukum than in Coriaria nepalensis. The weight loss pattern of these four species during decomposition could be best described by a double exponential decay model, and monthly decay rate coefficients (k) were in the order Q. fabri > C. nepalensis > I. cylindrica > P. aquilinum over a one-year period. Total nutrient (N, P and K) release of the four mulching materials ranged from 74.2 to 90.8% of the initial content after one year of decomposition, while nutrients were released in the order K > P>N. Nutrient release was fastest in C. nepalensis followed by Q. fabri, I. cylindrica, and slowest in P. aquilinum. Thus, taking into account initial nutrient content, nutrient additions and decay rates, the sprouts of woody species (Q. fabri and C. nepalensis) with massive foliage proved to be more important as nitrogen sources than the grass or fern.     

Soil fertility enhancement by planted tree-fallow species in the humid lowlands of Cameroon

Four shrub/tree species, Alchornea cordifolia, Pennisetum purpureum, Chromolaena odorata, and Calliandra calothyrsus were evaluated for their potential contribution to soil fertility restoration after two years fallow. Standing biomass, root distribution, nutrient content in the biomass, decomposition and nutrient release patterns, and association with mycorrhizae were the evaluation parameters. Alchornea and Pennisetum produced thehighest above-ground biomass, 66 t and 54 t/ha respectively. Pennisetum had more than 19 t/ha of root, 92% of which was in the 20 cm top soil. Alchornea had 74% of it roots in this soil layer, mostly as coarse roots while Calliandra had a deeper root system. Alchornea fallow accumulated more N and Ca, and Pennisetum fallow, more K than others, and mycorrhizae were mostly associated with Alchornea roots. The ranking of the different species for the decomposition rate was: Chromolaena > Pennisetum > Calliandra = Alchornea. Also release of nutrients during decomposition followed the order K > N > Ca. Alchornea and Pennisetum could be recommended as green manure species especially when high quantities of material are needed for weed or erosion control. Calliandra and Chromolaena, because of the flush of nutrient during early mass, loss can be used as mulch when the crop demand of nutrient is high. Alchornea decomposed slowly and therefore could be used to improve Chromolaena mulch, thus contributing to the build up of soil organic N and providing both short- and long-term nutrient release. This revised version was published online in June 2006 with corrections to the Cover Date.     

马尾松、桉树及其混交林土壤养分变化特征及评价

研究马尾松纯林、桉树纯林和3种不同模式混交林土壤养分含量的变化规律,并对其土壤养分状况进行主成分综合排序。结果表明：土壤有机质、全 N、全 P、水解 N、速效 P 和速效 K 皆表现为随土壤深度增加而减少,全 K 则表现相反;在各林分中,桉树纯林的土壤有机质、速效 P 含量最高,混交林次之。混交林则土壤全 N、全P 和速效 N 含量总体上大于纯林。马尾松纯林除在浅层土中速效 P 含量最高外,其他养分指标都低于桉树纯林和混交林。土壤养分综合评价为桉树纯林＞混交林 A＞混交林 C＞混交林 B＞马尾松纯林,表现为桉树纯林养分状况最好,马尾松纯林最差,与针阔树种的生长特性和凋落物养分归还能力有关。混交林中种植密度较大的混交林A 养分状况相对较好,表明其养分储存能力较好,消耗较少。而相同种植密度下,混交林 C 的林分土壤养分状况优于混交林 B。     

土壤微生物生物量及其影响因素研究

土壤微生物生物量的养分贮量比土壤有机质的小,但它既是植物所需养分的有效来源,还是养分的临时贮库。本文根据国内外研究结果综述了一些影响土壤微生物生物量动态的因素。这些因素主要包括 C 和 N 的限制、残留物和营养物质的经营、植物种类、土壤质地、土壤湿度和温度等。通过细致分析,我们认为未来研究应着重于探讨土地利用变化对热带和亚热带土壤微生物生物量的影响。参 55。     

Leaf-Litter Decomposition in an Oak-Conifer Forest in Himalaya: The Effects of Climate and Chemical Composition

The decomposition rates of certain dominant litter species inan undisturbed oak-conifer forest in the Kumaun Himalaya wereinvestigated by enclosing pre-weighed, newly senesced leavesin litter bags, placing these bags on the forest floor and determiningthe weight loss by recovering the bags at monthly intervals.Among the species examined, Daphne cannabina decomposed fastestand Cupressus torulosa decomposed most slowly. The former took6 months for complete decomposition, while Cupressus torulosadecomposed to about 72 per cent in 18 months. A linear combinationof rainfall and temperature explained 80 per cent of the variabilityin monthly weight loss. The rate of decomposition was also relatedto the initial chemical composition of the litter. Initially,several constituents showed a significant relation with decomposition;these are nitrogen, calcium, water soluble compounds, totalnonstructural carbohydrates, lignin, acid detergent cell wallcomponents and fibre. Of these, lignin and fibre contents showeda negative relation with weight loss while the others were positivelyrelated. As decomposition progressed, only nitrogen, lignin,ADCW and fibre content remained important. With further lapseof time only two constituents, nitrogen and lignin, remainedimportant. Among these the effect of nitrogen declined withtime while that of lignin increased. A linear combination ofnitrogen and lignin explained 59 per cent variability in theannual weight loss.     

秸秆覆盖对林地土壤肥力及树体生长的影响

秸秆覆盖降低了土壤温度,缓和了暴风雨对土壤的直接侵蚀,减少了水、土、肥的流失;促进了土壤有机质和营养物质的积累、分解和转化,从而复壮了树势,提高了树体的生长量及养分含量。本文通过综述秸秆覆盖的发展及其所产生的土壤效应、环境效应和对树体营养生长的影响,为探索秸秆覆盖对其它经济林的影响提供参考和依据。     

Litter and fine root dynamics of a relict sacred grove forest at Cherrapunji in north-eastern India

In a sacred grove climax forest (protected by the local Khasi tribe for religious reasons) at Cherrapunji in north-eastern India (mean annual rainfall, 10 372 mm), litter dynamics and related fine root dynamics were studied. Litterfall and its decomposition were very pronounced during the monsoon season, unlike in other rain forests in the region. High levels of nitrogen (2.13–3.58%) phosphorus (0.62–0.91%) and potassium (1.45–1.98%) in the leaf litter of four dominant tree species—Englehardtia spicata Bl., Echinocarpus dasycarpus Benth, Sysygium cuminii (L.) Skeels and Drimycarpus racemosus Hk.f.—suggest that these species help in conserving nutrient elements, thus ensuring their rapid recycling. Different species have different nutrient release patterns which are related to litter quality and seasonal environmental factors.

The highly developed fine root system (14 000 kg ha⁻¹) of which about 48% is located in the 0–10 cm soil depth, is important for nutrient storage and rapid recycling of nutrients. With a productivity of 3200 kg ha⁻¹ per year, the fine root component of the climax sacred grove ecosystem has a key role to play in tight nutrient cycling. It is concluded that these results are important in understanding the fragility of rainforest ecosystems and their management.     

西双版纳普文林场热带雨林恢复的实践

近50年来,通过树种调查、繁殖技术研究、造林试验,在西双版纳普文林场,成功地培育出8种热带雨林标志树种的人工林及7种热带山地雨林标志性树种的人工林,并散生培育热带雨林树种12个。还在热区退化地造林中评选出了2个造林先锋树种。它们既形成了高大整齐的林分外貌,也逐步形成了多层次多物种的森林复层结构。在这些人工林下出现了众多的雨林标志树种的天然更新幼苗幼树,说明这些人工林所形成的雨林环境具有可持续经营性。实践表明,西双版纳的热带雨林是可人工恢复的。     

Tree-grass relationships in open eucalypt woodlands of northeastern Australia: influence of trees on pasture productivity, forage quality and species distribution

Pasture yield, quality and species distributions were compared between zones around live and killed eucalypt trees at two woodland sites in northeast Queensland which differed markedly in soil fertility. Trees affected pasture quality and yield on an individual tree basis: N concentration and dry matter digestibility tended to be higher under trees than in inter-tree areas at both sites and pasture yields declined with distance from killed trees at the lower fertility site. However, the distribution of individual species did not vary markedly with distance from trees. Trees also affected pasture yield on a woodland basis: yields were greater where the trees were killed than under intact woodland. Soil under trees had higher levels of organic carbon and greater litter cover than soil in inter-tree areas. However, pasture yields did not generally reflect this fertility gradient since growth was limited by moisture availability due to drought conditions during the study period. Pasture N concentration was higher under trees that in inter-tree areas since plants under trees produced a similar amount of biomass as plants in inter-tree areas, but had access to higher nutrient levels. Trees appeared to have a greater effect on soil nutrient availability at the low fertility site. Live trees depressed pasture yields to a lesser degree at the low fertility site, demonstrating that the effects of trees on soil water availability (on a woodland scale) are less important when soil nutrients are more limiting to growth. These results indicate that, while removing trees may enhance pasture productivity, this benefit may be offset by a reduction in pasture quality. Given the beneficial effect of trees on soil nutrients, tree removal may also have longer term implications for soil nutrient dynamics. This revised version was published online in August 2006 with corrections to the Cover Date.     

海南霸王岭热带山地雨林林隙幼苗库动态规律研究

分析了海南岛霸王岭自然保护区热带山地雨林林隙内树种的种类组成、数量动态以及林隙年龄和林隙面积对树种组成及数量的影响。结果表明 :热带山地雨林林隙树种组成十分复杂 ,数量分布极不均匀 ,种间差异较大 ,2 7个林隙中共有乔木 116种 ,其中数量较多的 10个种的个体数量占到总个体数量的 4 7 0 3% ,而个体数量较少的 5 8个种的个体数仅占总个体数的 6 0 2 %。林隙面积大小对树种幼苗的种类组成和数量有较大影响 ,幼苗种密度和个体密度随林隙面积的增大而减小 ;幼树和成树在不同大小林隙中的种密度和个体密度变化不大。成熟林隙明显比早期林隙和中等年龄林隙有较大的幼苗个体密度和种密度 ,不同年龄林隙内幼树和成树的个体密度变化也不大。幼苗向幼树的转化率在大林隙和成熟林隙内明显高于小林隙和幼龄林隙。依据林隙树木幼苗数量变化的差异 ,可将林隙填充者分为 4种不同类型。