Long-term dynamics of vegetation and disturbance of a southern boreal spruce swamp forest

Abstract. Analysis of pollen, charcoal and loss-on-ignition in peat cores from a Picea aèies-dominated swamp forest in central Sweden show the vegetation changes and disturbance patterns over 9500 yr. Six major sequences of local vegetation development are identified: (A) Pinus period, ca. 9500–7000 cal. BP; (B) Open mire period (ca. 7000–4500 cal. BP; (C) Betula period, ca. 4500–2300 cal. BP; (D) Picea period (ca. 2300–1000 cal. BP; (E) Human impact period (ca. 1000–100 cal. BP); and (F) Period of human abandonment during the last ca. 100 yr. The swamp forest has been highly dynamic in response to various natural and anthropogenic disturbance agencies. Several fires have heavily influenced the vegetation development. During the last ca. 900 yr human influence has been important, initially from grazing and trampling by domesticated animals (ca. 1000–500 cal. BP), and subsequently small-scale cereal growing (ca. 400–100 BP). Cutting, burning and animal browsing influenced the structure and dynamics of the swamp forest by creating a more open stand and suppressing tree regeneration. Recent cessation of human impact has led to increased tree regeneration and a denser swamp forest stand. The present high biodiversity, and subsequent conservation interest does not result from long-term stability or absence of fire and human impact. However, in spite of repeated disturbances, a continuity of old and senescent trees produced a forest type with abundant dead wood. With the relatively minor importance of fire over long periods of time, the swamp forest developed a structure maintaining a high biological diversity. An important issue for maintaining long-term biodiversity in the boreal landscape must be to create a mosaic where different forest types are present, with a variety of structures, substrates and processes, to provide a certain degree of freedom for species to move around in the landscape.     

Analysing forest recovery after wildfire disturbance in boreal Siberia using remotely sensed vegetation indices

Wildfires have major effects on forest dynamics, succession and the carbon cycle in the boreal biome. They are a significant source of carbon emissions, and current observed changes in wildfire regimes due to changes in climate could affect the balance of the boreal carbon pool. A better understanding of postwildfire vegetation dynamics in boreal forests will help predict the future role of boreal forests as a carbon sink or source. Time series of Normalized Difference Vegetation Index (NDVI) and Normalized Difference Shortwave Infrared Index (NDSWIR) derived from Moderate Resolution Imaging Spectroradiometer (MODIS) aboard the Terra satellite were used to investigate whether characteristic temporal patterns exist for stands of different ages in the Siberian boreal forests and whether their postwildfire dynamics are influenced by variables such as prewildfire vegetation cover. Two types of forests, evergreen needle‐leaf (ENF) and deciduous needle‐leaf (DNF), were studied by analysing a sample of 78 burned forest areas. In order to study a longer time frame, a chronosequence of burned areas of different ages was built by coupling information on location and age provided by a forest burned area database (from 1992 to 2003) to MODIS NDVI and NDSWIR time series acquired from 2001 to 2005. For each of the burned areas, an adjacent unburned control plot representing the same forest type was selected, with the aim of separating the interannual variations caused by climate from changes in NDVI and NDSWIR behaviour due to a wildfire. The results suggest that it takes more than 13 years for the temporal NDVI and NDSWIR signal to recover fully after wildfire. NDSWIR, which is associated to canopy moisture, needs a longer recovery period than NDVI, which is associated to vegetation greenness. The results also suggest that variability observed in postwildfire NDVI and NDSWIR can be explained partially by the dominant forest type: while 13 years after a fire NDVI and NDSWIR are similar for ENF and DNF, the initial impact appears to be greater on the NDVI and NDSWIR of ENF, suggesting a faster recovery by ENF.     

林火干扰对北方针叶林林下植被的影响

林下植被在北方针叶林植被群落中的物种多样性最高, 且具有较高的生物量周转率和地上部分净初级生产力, 对北方针叶林生态系统功能起着重要作用。火干扰是决定北方针叶林林下植被结构与功能的一个重要景观过程。该文综述了火干扰是如何通过与地形、火前林冠组成的交互作用而影响环境资源和林下植被的。最近的研究表明: 林下植被能够影响火后树木更新苗的定植、重建速率及森林演替轨迹; 林下植被还会通过影响元素的生物地球化学过程(凋落物降解和养分循环)影响林下环境资源的数量与异质性。因此, 研究火后初期北方针叶林林下植被的动态变化, 对于物种多样性保护和森林管理具有重要意义。     

Spatial and temporal dynamics of Siberian silk moth large-scale outbreak in dark-needle coniferous tree stands in Altai

The spatial and temporal dynamics of fir stands damage caused by a large-scale outbreak of the Siberian silk moth (Dendrolimus sibiricus Tschetv.) in the Altai Mountains has been studied using remote sensing and GIS methods. It is ascertained that forest damages are dissimilar relative to the surface features. The pest outbreak initially occurred on the southwest slopes with mean values of 10° and elevation of 400 m asl. The damages further extended both upward and downward, involving slopes of high steepness and the eastern exposure. The total area of the dead stands comprised 6000 ha, 45% of which were lost due to secondary pest (xylophagous insects) attacks. It is indicated that the use of remote sensing made it possible to determine the beginning of a pest outbreak with one-decade precision. The Siberian silk moth large-scale outbreak occurred against an increase in air temperatures, a decrease in precipitation, and a reduction in late frosts. The tree plants weakened by the Siberian silk moth and water stress were affected by xylophagous insect attacks. The observed and predicted warming and climate aridity increase will facilitate Siberian silk moth outbreaks both within its range and northward of the current margins of the range.     

Sixty years of vegetation dynamics in a south boreal coniferous forest in southern Norway

Abstract. Vegetation data from permanent plots were collected in 1931, 1961 and 1991 in a south boreal forest 20 km north of Oslo in southern Norway. Major changes were found in the vegetation composition during those 60 years. The main changes were a reduction in the frequency of species and the frequency of joint occurrences of vascular species such as Andromeda polifolia, Calluna vulgaris, Cornus suecica, Eriophorum vaginatum, Maianthemum bifolium, Melampyrum pratense, Trientalis europaea, Vaccinium uliginosum and V. oxycoccus, and mosses, e.g. Dicranum fuscescens, Hylocomium splendens, Pleurozium schreberi, Ptilidium ciliare and Ptilium crista-castrensis. The observed changes were interpreted as being induced by internal processes e.g. notably a long-term change from paludified forest to mesic forest. In particular the growth of Picea abies seems to be a main driving force. The dominance of Picea abies and Vaccinium myrtillus appears to have rendered the conditions more unfavourable for other species. A doubling of the living stem biomass of P. abies during the last 67 yr shows that this old-growth forest has not yet reached a steady state. It was demonstrated that species such as Deschampsia flexuosa and Molinia caerulea did not increase in frequency in response to nitrogen deposition, as has occurred elsewhere in northern Europe. pH in the humus layer increased with 0.2 unit from 1961 to 1991. The results of this study indicate that protection from logging has initiated the reduction of species in the field layer and bottom layer. This study questions if monitoring of forest vegetation should be restricted to protected forests as is the practice in Scandinavia today. We recommend that also areas with some kind of selective cutting will be used for monitoring of forest vegetation.     

长白山暗针叶林苔藓植物生物量的研究

在长白山北坡暗针叶林对地面和树附生苔藓植物的生物量进行了测定.地面生苔藓采取样带调查取样法测定,树附生苔藓应用McCune方法对树干和树枝的附生苔藓生物量都做了细致的测定.结果表明,长白山暗针叶林中的苔藓植物分布很不均匀,随海拔变化差异很大,海拔1100m最低,仅为543kg·hm^-2;海拔1250m最高,达5097kg·hm^-2.苔藓植物生物量的变化对生境有很好的指示作用,特别是塔藓和拟垂枝藓的生物量随海拔的变化与森林系统的群落学特点有一定的相关性:在海拔1100～1700m,塔藓的生物量与臭冷杉的重要值变化趋势相近,随海拔升高而减少;拟垂枝藓的生物量与鱼鳞云杉重要值的变化趋势相似,随海拔升高而增加.此外,生物量随海拔的变化表明了不同苔藓植物对环境条件要求的差异,拟垂枝藓比塔藓水分条件要求更高.因此,生物量的研究在植物生理上也有一定的指示作用.     

长白山暗针叶林苔藓植物生物量的研究

在长白山北坡暗针叶林对地面和树附生苔藓植物的生物量进行了测定．地面生苔藓采取样带调查取样法测定,树附生苔藓应用McCune方法对树干和树枝的附生苔藓生物量都做了细致的测定．结果表明,长白山暗针叶林中的苔藓植物分布很不均匀,随海拔变化差异很大,海拔1100m最低,仅为543kg·hm^-2;海拔1250m最高,达5097kg·hm^-2．苔藓植物生物量的变化对生境有很好的指示作用,特别是塔藓和拟垂枝藓的生物量随海拔的变化与森林系统的群落学特点有一定的相关性：在海拔1100～1700m,塔藓的生物量与臭冷杉的重要值变化趋势相近,随海拔升高而减少;拟垂枝藓的生物量与鱼鳞云杉重要值的变化趋势相似,随海拔升高而增加．此外,生物量随海拔的变化表明了不同苔藓植物对环境条件要求的差异,拟垂枝藓比塔藓水分条件要求更高．因此,生物量的研究在植物生理上也有一定的指示作用．     

Soil organic sulfur dynamics in a coniferous forest

Sulfate microbial immobilization and the mineralization of organic S were measured in vitro in soil horizons (LFH, Ae, Bhf, Bf and C) of the Lake Laflamme watershed (47°17 N, 71°14 O) using ³⁵SO₄. LFH samples immobilized from 23 to 77% of the added ³⁵SO₄ within 2 to 11 days. The ³⁵SO₄ microbial immobilization increased with temperature and reached an asymptote after a few days. The mineral soil generally immobilized less than 20% of the added ³⁵SO₄, and an asymptote was reached after 2 days. An isotopic equilibrium was rapidly reached in mineral horizons. A two-compartment (SO₄ and organic S) model adequately described ³⁵SO₄ microbial immobilization kinetics. The active organic reservoir in the whole soil profile represented less than 1% of the total organic S. The average concentrations of dissolved organic S (DOS) in the soil solutions leaving the LFH, Bhf and Bf horizons were respectively 334, 282 and 143 µgL^–1. Assuming that the DOS decrease with soil depth corresponded to the quantities adsorbed in the B horizons, we estimated that 12 800 kgha^–1 of organic S could have been formed since the last glaciation, which is about 13 times the size of the actual B horizons reservoirs. Our results suggest that the organic S reservoirs present in mineral forest soils are mostly formed by the DOS adsorption resulting from incomplete litter decomposition in the humus layer. The capability of these horizons to immobilize SO₄ from the soil solution would be restricted to a 1% active fraction composed of microorganisms. Despite their refractory nature, these reservoirs can, however, be slowly decomposed by microorganisms and contribute to the S-SO₄ export from the watershed in the long term.     

Intensive ground vegetation growth mitigates the carbon loss after forest disturbance

Aims

Slow or failed tree regeneration after forest disturbance is increasingly observed in the central European Alps, potentially amplifying the carbon (C) loss from disturbance. We aimed at quantifying C dynamics of a poorly regenerating disturbance site with a special focus on the role of non-woody ground vegetation.

Methods

Soil CO₂ efflux, fine root biomass, ground vegetation biomass, tree increment and litter input were assessed in (i) an undisturbed section of a ~ 110 years old Norway spruce stand, (ii) in a disturbed section which was clear-cut six years ago (no tree regeneration), and (iii) in a disturbed section which was clear-cut three years ago (no tree regeneration).

Results

Total soil CO₂ efflux was similar across all stand sections (8.5 ± 0.2 to 8.9 ± 0.3 t C ha^?1 yr.^?1). The undisturbed forest served as atmospheric C sink (2.1 t C ha^?1 yr.^?1), whereas both clearings were C sources to the atmosphere. The source strength three years after disturbance (?5.5 t C ha^?1 yr.^?1) was almost twice as high as six years after disturbance (?2.9 t C ha^?1 yr.^?1), with declining heterotrophic soil respiration and the high productivity of dense graminoid ground vegetation mitigating C loss.

Conclusions

C loss after disturbance decreases with time and ground vegetation growth. Dense non-woody ground vegetation cover can hamper tree regeneration but simultaneously decrease the ecosystem C loss. The role of ground vegetation should be more explicitly taken into account in forest C budgets assessing disturbance effects.

     

卧龙自然保护区亚高山暗针叶林林隙特征研究

以卧龙亚高山暗针叶林为研究对象,分析了森林群落林隙的大小结构、形成方式及林隙形成木(GM)的结构特征．结果表明,林隙密度为18．5个·hm^-2冠林隙和扩展林隙分别占森林面积的28．4％和60．0％;冠林隙的大小变化在10～1134．7m^2之间,平均面积为153．45m^2;扩展林隙的大小变化在84．11～1646．3m^2之间,平均面积为324．34m^2;平均每个林隙的形成木为5．14株,单株形成木形成的林隙只占8．1％．不同形成木类型对林隙形成的贡献大小次序为：折干>掘根>枯立>断梢;暗针叶林林隙大多由岷江冷杉、铁杉、糙皮桦形成．径级在60～70cm,高度在30～35m之间的林冠上层的岷江冷杉,发生折倒形成林隙的可能性最大．     

Performance of the potentially invasive Siberian moth Dendrolimus superans sibiricus on coniferous species in Europe

1 The native range of the Siberian moth extends from the Pacific Ocean (Russian Far East, Japan and Northern Korea) across Siberia, Northern China and Mongolia to the Ural Mountains. At the beginning of the 21st Century, this species was documented west of the Ural Mountains in the Republic of Mari El, indicating range extension toward the west.
2 The Siberian moth has recently been suggested for regulation as a quarantine pest for European and Mediterranean Plant Protection Organization member countries. However, no specific report on European host plants for this pest has been published so far.
3 In the present study, larval host plant choice and performance was tested for the first time on coniferous tree species that are widely distributed and of commercial value in Europe.
4 Based on dual-choice tests on neonates and mortality, developmental duration and relative growth rates of the first- to sixth-instar larvae, we found European larch Larix decidua to be the most suitable host for the moth larvae, whereas European black pine Pinus nigra and Scots pine Pinus sylvestris were the poorest hosts. The remaining conifer species tested, European silver fir Abies alba , Nordmann fir Abies nordmanniana , and Norway spruce Picea abies , were intermediate host plants. Douglas-fir Pseudotsuga menziesii , originating from North America, was chosen by the larvae to the same extend as European larch, and was also highly suitable for larval development.
5 If the moth is introduced to European countries, it will become damaging in stands of European larch and Douglas-fir, mixed stands of fir and spruce; however, it will be less damaging in forests dominated by two-needle pines.
6 We predict that Dendrolimus superans sibiricus will be able to survive and develop on the main European coniferous tree species, including non-native coniferous tree species, resulting in severe damage to large areas of forests.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

西藏色季拉山暗针叶林凋落物层化学性质研究

The storage and chemical properties of the forest litter in dark coniferous forest of Sejila Mountain were studied. The results showed that the existing storage was 5. 863t·hm^-2 and the annual litter fall was 0. 3205 t·hm^-2 It implied that the forest litter decomposed slowly and accumulated quickly, and the turnover of nutrient circles was slow. The contents of N, Ca, Na, and Mn nutrient elements in litter layer were in the order of un-decomposed layer (U layer) > semi-decomposed layer (S layer) > decomposed layer (D layer), those of K, Fe, and Mg were in the order of D layer > S layer > U layer, and P element content was in the order of U layer > D layer> S layer. The pool of elements was 78. 483 kg·hm^-2 N, 3. 843 kg·hm^-2P, 48. 205 kg·hm^-2 K, 23.115 kg·hm^-2 Ca, 13. 157 kg·hm^-2 Na, 30.554 kg·hm^-2 Fe, 2. 113 kg·hm^-2 Mn and 27. 513 kg·hm^-2 Mg. The turnover of forest litter was the total of nutrient release accumulation. K, Fe, and Mg were enriched, and N,Ca, Na, Mn, and P were released with the turnover rate in the order of N > Ca > Na > Mn >P.     

长白山暗针叶林苔藓植物在养分循环中的作用

测定了长白山暗针叶林5种主要苔藓植物养分含量,并结合生物量的测定结果,推算单位面积的苔藓植物养分含量.结果表明,单位面积苔藓植物氮含量为5.371 kg·hm^-2,是乔、灌、草总氮量的12.22%;钾含量为12.02 kg·hm^-2,是乔、灌、草总钾量的5.63%;磷含量为31.679 kg·hm^-2,是乔、灌、草总磷量的70.57%,超过了乔木磷含量,是草本植物磷含量的2.倍、灌木磷含量的792倍,苔藓植物是暗针叶林重要的磷库.同时测定不同苔藓盖度下土壤中的养分含量.方差分析结果显示,不同苔藓盖度下土壤中氮、钾含量差别不大,磷含量差异显著,没有苔藓覆盖的土壤中磷含量最高,为0.19±0.023 g·kg^-1;苔藓盖度为第6级(80%～100%),土壤中磷含量最低,为0.36±0.017 g·kg^-1,随苔藓盖度的增加土壤中磷含量有减少的趋势.     

长白山北坡暗针叶林群落特征

长白山北坡的暗针叶林带分布海拔为１１００～１８５０ｍ,在野外调查的基础上,对暗针叶林沿海拔梯度物种组成的分布格局进行了研究。文中用α和β多样性指数来度量森林群落的多样性,用群落系数（本质上是一种β多样性指数）来度量不同群落或样地间的相似性,同时用欧氏距离和Ｗａｒｄ法对所有样地进行了聚类分析。研究结果表明主要树种云杉和冷杉在整个暗针叶林带均分布,而伴生树种和多数灌木出现于一定海拔范围内;沿海拔梯度物     

Long-term effects of clear-felling on vegetation dynamics and species diversity in a boreal pine forest

The understorey vegetation in a lichen–Scots pine forest was monitored during 20 years before and after clear-felling. Plots with and without logging residues were compared concerning the general pattern of the vegetation dynamics and changes in species composition, dominance, richness, evenness and diversity. The succession of both treatments had a clear principal component analysis (PCA) pattern of a 'stepwise arch-shaped diverging' trend mainly driven by 'pioneer' lichens, 'reindeer' lichens and Calluna vulgaris. The difference between the residue treatments was significant regarding succession of vascular plants, bryophytes and 'reindeer' lichens. The nitrogen indicators Epilobium angustifolium and Deschampsia flexuosa were favoured on plots with logging residues.     

Occupancy of red‐naped sapsuckers in a coniferous forest: using LiDAR to understand effects of vegetation structure and disturbance

Red‐naped sapsuckers (Sphyrapicus nuchalis) are functionally important because they create sapwells and cavities that other species use for food and nesting. Red‐naped sapsucker ecology within aspen (Populus tremuloides) has been well studied, but relatively little is known about red‐naped sapsuckers in conifer forests. We used light detection and ranging (LiDAR) data to examine occupancy patterns of red‐naped sapsuckers in a conifer‐dominated system. We surveyed for sapsuckers at 162 sites in northern Idaho, USA, during 2009 and 2010. We used occupancy models and an information‐theoretic approach to model sapsucker occupancy as a function of four LiDAR‐based metrics that characterized vegetation structure and tree harvest, and one non‐LiDAR metric that characterized distance to major roads. We evaluated model support across a range of territory sizes using Akaike's information criterion. Top model support was highest at the 4‐ha extent, which suggested that 4 ha was the most relevant scale describing sapsucker occupancy. Sapsuckers were positively associated with variation of canopy height and harvested area, and negatively associated with shrub and large tree density. These results suggest that harvest regimes and structural diversity of vegetation at moderate extents (e.g., 4 ha) largely influence occurrence of red‐naped sapsuckers in conifer forests. Given the current and projected declines of aspen populations, it will be increasingly important to assess habitat relationships, as well as demographic characteristics, of aspen‐associated species such as red‐naped sapsuckers within conifer‐dominated systems to meet future management and conservation goals.     

Modelling carbon dynamics in coniferous forest soils in a temperature gradient

Climate change and changes in land use will alter the stores of carbon and turnover of soil organic matter. We have used a theory for carbon cycles in terrestrial ecosystems to analyse changes in soil organic matter turnover in coniferous forests. The central concepts of the theory are a continuously changing substrate quality, a constant decomposer efficiency and a climatically controlled decomposer growth rate. Measurements on litter production and soil carbon stores from field experiments have been used to successfully validate the model predictions. Measured litter production increased with increasing temperature but the response was not identical for forests of different vegetation types which reflect variations in productivity. The temperature response of needle-litter production and decomposition rate were strongest in the most productive forests and weakest for the low productive forests. Initial decay rates of soil C store from steady state showed the same trend in temperature response as decay of a single litter cohort did, but the absolute values are 16% of the decay rates of a single litter cohort. Predicted soil C ranged from 5 to 9 kg C m^–2. There exists a remarkable variation in forest soil C store response to temperature; the magnitude and even the sign depends on productivity as defined by vegetation type. The assumption that, in general, decomposition rates increase more than NPP with temperature, and consequently, soil C stores should decrease in response to a climate warming, seems therefore too simplistic.     

Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance

Decomposition of typhoon-generated and normal leaf litter and their release patterns for eight nutrient elements were investigated over 3 yr using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. Two common tree species, Castanopsis sieboldii and Schima wallichii, representative of the vegetation and differing in their foliar traits, were selected. The elements analyzed were N, P, K, Ca, Mg, Na, Al, Fe and Mn. Dry mass loss at the end of study varied in the order: typhoon green leaves > typhoon yellow leaves > normal leaves falling for both species. For the same litter type, Schima decomposed faster than Castanopsis. Dry mass remaining after 2 yr of decomposition was positively correlated with initial C:N and C:P ratios. There was a wide range in patterns of nutrient concentration, from a net accumulation to a rapid loss in decomposition. Leaf litter generated by typhoons decomposed more rapidly than did the normal litter, with rapid losses for N and P. Analysis of initial quality for the different litter types showed that the C:P ratios were extremely high (range 896 – 2467) but the P:N ratios were < 0.05 (range 0.02 – 0.04), indicating a likely P-limitation for this forest. On average 32% less N and 60% less P was retranslocated from the typhoon-generated green leaves than from the normal litter for the two species, Castanopsis and Schima. An estimated 2.13 g m^–2 yr^–1 more N and 0.07 g m^–2 yr^–1 more P was transferred to the soil as result of typhoon disturbances, which were as high as 52% of N and 74% of P inputted from leaf litter annually in a normal year. Typhoon-driven maintenance of rapid P cycling appears to be an important mechanism by which growth of this Okinawan subtropical forest is maintained.