南亚热带森林不同林型功能性状分布格局及其驱动机制

植物功能性状能反映植物对环境变化的响应,研究植物功能性状的分布格局有助于揭示群落的构建过程及其内在作用机制。该研究以鼎湖山南亚热带山地常绿阔叶林和沟谷雨林为研究对象,采集并测量了样地中木本植物的12种不同的功能性状,分别以5 m×5 m、10 m×10 m、20 m×20 m的样方为尺度单元,通过计算平均成对性状距离指数来探讨群落中功能性状的分布格局及其驱动机制。结果表明,两个林型的群落中12个功能性状均存在不同程度变异,但功能性状在群落间的差异不显著(P>0.05)。两个林型的群落中功能性状空间分布格局均具有尺度依赖性,但不同尺度的驱动机制有差异,随着空间尺度的增大,山地常绿阔叶林的功能性状空间分布格局主要驱动机制由环境过滤转为扩散限制;沟谷雨林的由环境过滤和相似性限制转为扩散限制,两个林型在20m×20m空间尺度上都是扩散限制。生态位分化和扩散限制综合作用于鼎湖山南亚热带山地常绿阔叶林和沟谷雨林的群落功能性状分布格局的产生及其群落构建过程,二者的贡献作用会随空间尺度发生变化。坡度是影响山地常绿阔叶林功能性状分布格局的最关键地形因子,海拔是影响沟谷雨林的最关键地形因子。     

Borreliae in Ixodes ricinus ticks feeding on humans

A total of 298 Ixodes ricinus (L.) ticks (Acari: Ixodidae) feeding on humans in the Czech Republic were tested for borreliae (Borrelia burgdorferi sensu lato) by darkfield microscopy between 1997 and 2003. A majority (68%) of the supplied I. ricinus ticks were nymphs, 25% were females and 7% were larvae. Overall, 20% of 74 examined females and 9% of 203 examined nymphs (but none of 21 examined larvae) were infected with borreliae. The proportion of ticks with a high infection load (>100 spirochetes) was 4% in females and 2% in nymphal I. ricinus. During the year, the highest numbers and proportions of infected nymphal and female ticks were taken from humans in June. Detection of borreliae in the ticks feeding on humans might be helpful in the prophylaxis of Lyme borreliosis.     

Effects of canopy gaps on the genetic structure of Camellia japonica saplings in a Japanese old-growth evergreen forest

The genetic structure of Camellia japonica saplings was investigated in relation to canopy conditions in an old-growth evergreen forest in Tsushima, Japan. To elucidate effects of canopy gaps on genetic structure, a 1 ha study site was divided into 20 x 20 m quadrats, which were classified into a gap quadrats (GAP), closed canopy quadrats (CLS) and mixed quadrats. Five GAP quadrats and six CLS quadrats were analyzed separately. Isolation-by-distance was tested by examining the correlation between genetic distance and geographic distance. A significant positive correlation was detected for GAP quadrats, whilst that for CLS quadrats was significantly smaller and not significantly different from zero. On the other hand, an analysis using Moran's I spatial autocorrelation coefficients indicates that the genetic structure is weaker in GAP quadrats than in CLS quadrats in short distance classes. The values were significantly positive for both types of quadrat. These results, along with our field observations on flowering, suggest that canopy gaps affect the genetic structure of C. japonica saplings in two distinct ways. First, canopy gaps may promote flowering and mating in an isolation-by-distance manner within canopy gaps. Second, canopy gaps may promote seed production and resulting overlap in seed shadows may weaken fine-scale genetic structures.     

SPATIAL PATTERNING AND COMPETITION IN AN ASPEN-WHITE PINE SUCCESSIONAL SYSTEM

The spatial patterning of and competition between established large-toothed aspen (Populus grandidentata) and invading white pine (Pinus strobus) were examined in a 65-yr-old forest system at the AuSable Trails Institute of Environmental Studies, Kalkaska County, Michigan. Density, diameter, and distance measures were recorded for tree species on two 40 × 40 m grids containing 64 quadrats each 5 × 5 m. Several indices of dispersion were used to assess pattern and regression analysis of nearest neighbor distance to tree diameter provided an index of competition. The results indicate strong aggregation for both aspen and pine on a large scale, but regularity of pattern for pines in relation to their nearest neighbor. This regularity appears to be due to competition with established aspens and suggests that the spatial patterning of the aspens acts to control the spacing and subsequent development of the invading white pines.     

Use of CFSE staining of borreliae in studies on the interaction between borreliae and human neutrophils

Background  

Species of the tick-transmitted spirochete group Borrelia burgdorferi sensu lato (B. burgdorferi) cause Lyme borreliosis. Acute borrelial infection of the skin has unusual characteristics with only a mild local inflammatory response suggesting that the interaction between borreliae and the cells of the first-line defence might differ from that of other bacteria. It has been reported that human neutrophils phagocytose motile borreliae through an unconventional mechanism (tube phagocytosis) which is not observed with non-motile borreliae. Therefore, it would be of great interest to visualise the bacteria by a method not affecting motility and viability of borreliae to be able to study their interaction with the cells of the innate immunity. Carboxyfluorescein diacetate, succinimidyl ester (CFSE) labelling has been previously used for studying the adhesion of labelled bacteria to host cells and the uptake of labelled substrates by various cells using flow cytometry.     

山西高原草地景观的数量分类与排序

草地植被空间格局研究是草地景观研究中一项重要的基础工作。空间格局研究不仅限于描述景观的空间特征,而是要深入分析空间格局的分布规律及影响格局形成的原因及机制。以1∶50万《山西省草地类型图》、《山西省行政区域图》及《山西省地形图》为主要信息源,借助GIS技术,应用TWINSPAN和DCA对山西高原草地景观的1127个样方进行研究。结果表明:(1)在TWINSPAN分类结果中,31个群丛组从左到右的排列顺序很好地揭示了山西高原草地景观的纬向地带性分布规律。即从40°43'N向34°34'N依次分布着北部草原、中部灌草丛和草丛及南部的暖温带草丛。(2)1127个样方DCA排序图纵轴揭示了明显的海拔梯度,即由上至下海拔逐渐降低,相应地草地景观变化为亚高山草甸、灌草丛、草原及河漫滩草甸;DCA排序图的横轴揭示了明显的水分梯度,即从左到右水分逐渐增加。两大优势群丛组样方及44个优势群丛组物种DCA的排序图都很好地刻画了山西高原水热组合的变化,即从西北向东南由于地形地貌引起的水热组合由冷干向暖湿逐渐变化,水热组合的梯度变化进一步影响了草地景观整体的空间分布格局。(3)GIS技术与数量生态学方法相结合有助于丰富景观生态学研究的技术体系。     

Takahe Valley Hut: a focal point for weed invasion in an isolated area of Fiordland National Park, New Zealand

The role of backcountry huts as focal points for weed establishment and spread into New Zealand’s national parks has received little attention. In this study we describe the pattern of weed spread around Takahe Valley Hut, Murchison Mountains, Fiordland National Park. Established in 1948, the hut is located at 900 m a.s.l. at the ecotone between Nothofagus forest and valley floor shrubland/grassland. We recorded the distribution of vascular plants in quadrats (110) placed by restricted randomisation around the hut, and measured relative irradiance and distance from the hut. Nine exotic species, mostly grasses, were recorded, the most frequent being Agrostis capillaris (34%). The majority of occurrences of exotic plant species were located in the immediate vicinity (less than 5 m) of the hut but two exotic species (Agrostis capillaris and Dactylis glomerata) ranged more widely. Exotic species were present in well-drained shrubland and grassland but did not extend far into Nothofagus forest or onto infertile wetlands. The percentage of exotic species in quadrats declined significantly with distance from the hut. There was no linear relationship between the percentage of exotic species and relative irradiance. When forest quadrats were excluded, the number of native species in quadrats was negatively correlated with the number of exotic species, suggesting competitive displacement of native species by exotics in non-forest habitats. Long-term persistence of most exotic species at this site depends on physical disturbance and nutrient enrichment associated with human activities at the hut site. However the maintenance of this species pool has provided sufficient propagule pressure for some exotic species to disperse into the wider area. Weed accumulation around huts can be reduced by locating huts in vegetation types that are more resistant to invasion, and maintaining facilities to eliminate local weed infestations.     

木论喀斯特峰丛洼地森林群落空间格局及环境解释

基于广西壮族自治区木论国家级自然保护区典型峰丛洼地景观尺度内不同微生境条件和植物群落类型50个样地 (20 m × 20 m)的系统取样调查, 用二元物种指示方法(TWINSPAN)对样地内胸径(DBH) ≥ 1 cm的木本植物进行分类, 选择10个土壤环境因子和5个空间因子, 利用除趋势典范对应分析(DCCA)研究了森林群落分布的土壤环境与空间格局, 并给予定量化的合理解释。结果如下: 1) TWINSPAN将森林群落划分为11组, 在三级水平上分为4类生态群落类型。2) DCCA第一排序轴集中了排序的大部分信息, 突出反映了各森林群落所在的坡向和土壤主要养分梯度, 沿第一轴从左到右, 坡向由阴转阳, 岩石裸露率越来越高, 土壤主要养分逐渐降低, 森林群落分别出现了由原生性和耐阴性强逐步向阳性先锋树种为主的次生林和人工林变化的格局。3)因子分离分析结果表明, 土壤环境因子对森林群落分布格局的解释能力为39.16%, 其中21.02%单纯由土壤环境因子所引起, 空间因子的解释能力为31.34%, 其中13.16%独立于土壤环境的变化, 18.15%是土壤环境和空间因子相互耦合作用的结果, 不可解释部分达47.66%, 表明喀斯特峰丛洼地森林群落的物种共存受生态位分化理论和中性理论双重控制。     

Effect of forest clearing on the abundance of Ixodes ricinus ticks and the prevalence of Borrelia burgdorferi s.l

Questing Ixodes ricinus L. (Acari: Ixodidae) ticks were collected on a forest trail that had been completely cleared of shrubs and ground vegetation in winter 2002 and on a nearby control uncleared forest transect in South Moravia (Czech Republic). Samples were collected each May in 2003, 2004 and 2005. Nymphal ticks were 3.4 times, 1.9 times and 1.2 times less frequent on cleared forest than on uncleared forest trails in the three respective years, whereas adult tick abundance was 27.2 times, 4.0 times and 2.2 times lower, respectively. The ticks were examined for borreliae by dark-field microscopy: prevalence of nymphal ticks infected with Borrelia burgdorferi sensu lato (12.6% to 20.0%) did not differ significantly between the cleared and uncleared trail during the 3 years. In conclusion, the habitat modification appeared to result in a decreased abundance of I. ricinus as well as a reduced frequency of infected ticks (and thus indirectly a lower potential risk of Lyme borreliosis), which lasted, however, for only 2 years. Eight cultures of borreliae isolated from the ticks were all identified as the 'ornithophilic' genomic species Borrelia garinii, possibly indicating a greater role of forest birds than that of forest rodents as the hosts of immature I. ricinus in the tick (and borrelial) colonization of the cleared part of the forest.     

Heterogeneity of terrestrial bromeliad colonies and regeneration of Acacia praecox (Fabaceae) in a humid-subtropical-Chaco forest, Argentina

In several tropical and subtropical forests, plants of the understorey act as an ecological filter that differentially affects woody species regeneration. In convex sectors of the Schinopsis balansae (Anacardiaceae) forests of the Southeastern Chaco there are dense colonies of terrestrial bromeliads. These may influence forest regeneration by intercepting rain water and propagules in their tanks. Within colonies, the spatial distribution of bromeliads is clumped because their clonal growth leaves numerous internal gaps. In this study we describe the internal heterogeneity of three bromeliad colonies (plots) and analyze how this heterogeneity affects Acacia praecox regeneration (i.e. seedling recruitment and survival). In January 1996, we randomly placed three transects with 150 contiguous quadrats of 100 cm(2) in each plot. For each quadrat we recorded the type of floor cover (i.e. bromeliads, herbs, litter, or bare soil) and the presence of A. praecox seeds or seedlings. In July 1996 we relocated the transects and recorded seedling survival. Bromeliad colonies showed a high internal heterogeneity. Almost half of the 450 quadrats were covered by two terrestrial bromeliads. Aechmea distichantha was recorded in 81% of all quadrats with bromeliads, and Bromelia serra in the others. All quadrats with bromeliads were covered by litter. Half of them were occupied by the bases of bromeliads and the others were covered by their leaves. In contrast, where bromeliads were not present, soil surface was covered by litter in 83% and by herbaceous vegetation in 11% of the quadrats; very few quadrats were covered by bare soil. In January 1996, we recorded 127 seeds and 176 seedlings of A. praecox. Seed and seedling densities of A. praecox were similar in quadrats with and without bromeliads, but variability in seedling density of A. praecox was higher within than among plots. Seed density was higher in quadrats covered by bromeliad leaves than inside the tanks. Seedling survival of A. praecox was slightly higher in quadrats with bromeliads in only one of the three plots. No seedling survived inside the bromeliad tanks. Apparently. bromeliad colonies have no effect on seedling regeneration of A. praecox.     

Functional diversity versus species diversity: relationships with habitat heterogeneity at multiple scales in a subtropical evergreen broad-leaved forest

Understanding the relationship between functional and species diversity as well as their association with habitat heterogeneity can help reveal the mechanisms of species coexistence in ecological communities. However, these interactions have been poorly studied in subtropical forests. In this paper, we evaluated functional diversity (as measured by Rao’s Q) and traditional species diversity (based on Simpson’s index) in a 24 ha forest plot in a subtropical evergreen broad-leaved forest (EBLF) in China. We compared the sensitivities of functional and species diversity to topographic variables (elevation, convexity, slope and aspect) at multiple spatial scales based on 10 × 10, 20 × 20, 40 × 40 and 50 × 50 m quadrats. Functional and species diversity were found to have different distribution patterns along a topographical gradient, with functional diversity better explained by topography than was species diversity using a spatial autocorrelation regression error model. Furthermore, functional diversity had a significantly greater association with topographic variables than species diversity in both adult and young trees; in both cases, the strength of the diversity-habitat association increased with quadrat size. We conclude that functional diversity reflects a greater diversity-habitat association in EBLF than does species diversity, and that the association depends on the spatial scale and life stages of the woody plants under evaluation.     

Species–topography association in a species-rich subtropical forest of China

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m²) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.     

A Study on the Structural Dynamics and Spacial Pattern of Vatica xishuangbannaensis Population

The contiguous grid quadrats were applied to sampling for field data and the static life table of Vatica Xishuangbannaensis population, which is one of main dominant and rare species of tropical seasonal rain forest was set up, in Bubeng region situated in the south of Yunnan province (about N 20°30′,E 101°35′ ). The population was an unstable intermittent population. The spatial pattern of the population was of the contagious distribution-random distribution-uniform distribution from sapling to adult tree. The type of the population regeneration obeys the mosaic or cyclical theory of regeneration. According to population dynamics and spacial pattern the authors have proposed advancal method of managenment and administration, and ways of protection and utilization of V. xishuangbannaensis population.     

Habitat niche specialization in an understory species in a warm temperate forest

Relationships between microhabitat variables; understory light conditions in summer and winter, altitude, slope inclination and topographic categories (valley, ridge, and slope) and the distribution of Aucuba japonica Thunb. (Cornaceae), a common understory shrub species in Japan were examined using non-contagious 66, 20 × 20 m² quadrats. The Morishita’s I _δ suggested that A. japonica distributions were strongly heterogeneous among the quadrats. Therefore positive spatial autocorrelation of A. japonica at a within-quadrat level (≤20 m) was obvious. Moran’s I statistics showed a significant positive spatial autocorrelation in A. japonica abundance within the distance shorter than 60 m. But the partial Mantel tests suggested that the mass effect from neighboring quadrats would little explain A. japonica abundance in a quadrat. The partial Mantel tests also clearly showed that A. japonica distributions were strongly structured by topography and understory light conditions. Using Monte Carlo randomization tests, we found that A. japonica was aggregately distributed in quadrats in valley which were covered by deciduous canopies. Better understory light conditions in winter together with valley edaphic conditions may increase the abundance of A. japonica there. It is concluded that habitat niche specialization is important in structuring distribution of A. japonica in this forest community.     

黄土丘陵沟壑区坡面尺度土壤水分空间变异及影响因子

土壤水分空间分布特征及其影响因子是土壤前期含水量模拟和小流域产流机制研究的重要内容,也是半干旱地区进行生态建设的重要参考。通过对黄土高原典型坡面雨季前后100 cm深度内土壤含水量进行观测,分析地形、植被和雨季对土壤水分空间分布的影响。基本统计分析显示,土壤水分的空间异质性在上层(<20 cm)较小,在下层(>40 cm)较大。坡面尺度上,土壤含水量的空间差异主要表现在不同植被类型之间,而不是坡位之间。各覆被类型的土壤含水量相对大小为荒草地>8年生刺槐林>20年生刺槐林>沙棘林。即使沙棘林和刺槐林位于更利于获取土壤水分的地形条件下,其土壤含水量仍然明显低于荒草地。地形对土壤水分的影响被植被类型的影响所掩盖。上述规律在雨季前后都有明显表现。因此,完全基于地形指数的土壤水分预测模型在黄土高原应该慎用,植被类型应该作为土壤水分空间预测的一个重要参数。雨季使土壤含水量整体提高,但是土壤水分空间分布格局并没有根本改变,高处仍高,低处仍低,各样点处的土壤含水量在雨季前后达到显著相关水平,说明土壤水分空间格局并不是瞬时状态,而具有明显的时间稳定性。     

Vegetation development after the exclusion of grazing cattle in a meadow area in the south of Sweden

A study of 24 years of secondary succession was based on data from semi-permanent quadrats from investigations in 1952, 1955, 1968 and 1976, involving two exclosures, the first in a meadow grazed for hundreds of years but now abandoned, the second in a meadow mowed for hundreds of years and grazed for the last 50 years. A first order classification of quadrats produced units, which formed distinct spatial patterns indicating similar gradients, but also differences in response to the ceased grazing, in the two exclosures. A second order classification of units into groups revealed a rather simple structure of spatial and temporal relations. Eleven groups of species with similar behaviour could be recognized within a system of spatial and temporal species distributions. The vegetation in both exclosures developed towards an increased differentiation and heterogeneity or patchiness. The border between two soil types was clearly reflected in the spatial pattern of units. Rates of change were greatest in the beginning and were shown to closely follow logarithmic functions of time. The average number of species per m² decreased in all plots, in some cases as much as 50%. It was concluded that the diversity had decreased as a consequence of decreased species richness, decreased evenness and decreased pattern diversity. Many individual species distributions showed a pattern of nuclei surrounded with marginal belts. Differences in rate of change and persistence of spatial patterns between the plots, could be attributed to the differences in management history. These differences have disappeared as the succession has proceeded. The two exclosures have instead both conformed to the same floristic gradient, in turn based on a similar pattern of environmental gradients, primarily moisture. In the observed changes the emphasis was on the shifting importance of competing species populations, as some gained in importance at the expense of others. Competition has so far been a more important process in the vegetation development than immigration/extinction rates.Nomenclature follows Lid (1974) for vascular plants and Nyholm (1954–69) for mosses.This paper is the third part of a doctor's thesis from the Department of Plant Ecology, University of Lund. I wish to thank Prof. Nils Malmer for valuable advice and discussions, Mrs Mimmi Varga for drawing the figures and all my colleagues at the Department for their help and support.     

不同密度条件下芨芨草空间格局对环境胁迫的响应

基于小尺度上植物间相互作用与空间格局的高度相关性,选用可避免环境异质性影响的K2点格局函数,研究了沙枣-芨芨草群落中63个芨芨草样方(3密度条件×3生境条件×7重复)中芨芨草种群在小尺度下(0—0.5 m)的空间格局,探讨了在3种密度条件(高、中、低)及3种生境条件下(冠盖区、过渡区、空旷区),芨芨草种群空间格局对土壤理化性质胁迫(盐分、有机质及容重)程度的响应。研究结果表明,就土壤盐分,养分含量及土壤容重而言,土壤理化性质胁迫程度沿冠盖区、过渡区和空旷区增加的趋势。相应地,芨芨草种群呈聚集分布的样方比例在沿冠盖区(6/21)较低,而在过渡区(11/21)和空旷区(11/21)较高。然而,在不同密度条件下,芨芨草种群空间格局对土壤性质胁迫的响应不同。在低密度条件下,芨芨草种群在冠盖区多数为聚集分布(4/7),在过渡区和空旷区全部为聚集分布;中密度条件下,芨芨草聚集分布样方比例沿冠盖区-过渡区-空旷区梯度增加(分别为2/7,3/7,4/7),但低于同等胁迫条件下低密度样方中聚集分布数量;高密度条件下,在各胁迫条件下,除了一个过渡区高密度样方,其余芨芨草种群均为随机分布。总体上,随着环境胁迫增强,在中低密度下,芨芨草种群趋向于聚集分布;但在高密度下,芨芨草种群均以随机分布为主。此外,芨芨草种群空间格局随密度变化趋势比随土壤理化性质胁迫梯度变化趋势更加明显,可能表明相对于土壤理化性质胁迫程度,芨芨草种群密度对其空间格局影响更大。因此,在考虑芨芨草种群空间格局对环境胁迫的响应时,应当考虑种群密度因素。     

Phylogenetic relatedness explains highly interconnected and nested symbiotic networks of woody plants and arbuscular mycorrhizal fungi in a Chinese subtropical forest

Elucidating symbiotic relationships between arbuscular mycorrhizal fungi (AMF) and plants contributes to a better understanding of their reciprocally dependent coexistence and community assembly. However, the main drivers of plant and AMF community assembly remain unclear. In this study, we examined AMF communities from 166 root samples of 17 woody plant species from 10 quadrats in a Chinese subtropical forest using 454 pyrosequencing of 18S rRNA gene to describe symbiotic AMF–plant association. Our results show the woody plant–AMF networks to be highly interconnected and nested, but in antimodular and antispecialized manners. The nonrandom pattern in the woody plant–AMF network was explained by plant and AMF phylogenies, with a tendency for a stronger phylogenetic signal by plant than AMF phylogeny. This study suggests that the phylogenetic niche conservatism in woody plants and their AMF symbionts could contribute to interdependent AMF and plant community assembly in this subtropical forest ecosystem.     

南亚热带森林种群分布格局取样技术研究

森林群落中植物种群分布格局测定和分析的结果,密切依赖于野外取样的技术手段及资料的可靠性。本文通过鼎湖山自然保护区三个不同森林群落类型的取样,比较了随机样方、相邻格子样方、中心点四分法、最近相邻法和最近个体法在野外操作、数据整理以及种群分布格局测定分析中的效用,对各方法在南亚热带森林群落中进行种群分布格局研究中的适用性予以客观评价。研究表明,同一样地中不同取样方法和测定方法会导致不同结果。随机样方取样数据的测定结果受样方大小影响,而几种无样地方法在种类复杂的群落中,应用受到限制。相邻格子样方适用于各群落类型中种群分布格局的测定与分析。     

Multiple canopy opening effects on recruited saplings in a typhoon-disturbed tropical rainforest,Taiwan

To investigate the influence of multiple canopy openings on the composition and diversity of recruited saplings in a forest frequently disturbed by typhoons. We conducted tree-by-tree censuses (diameter at breast height ≥ 1 cm) and mapped gaps (canopy height < 5 m) in 1993, 2000, 2008, and 2013 in a tropical mountain zonal foothill evergreen broad-leaved forest in Nanjenshan Nature Reserve, Taiwan. We analyzed the composition and diversity of recruited saplings within a 2.1 ha plot (840 sampling quadrats (5 m × 5 m)) with variable numbers of canopy openings recorded during the study period. Composition of recruited saplings was dissimilar between quadrats that stayed opened and those that stayed closed throughout the study period (pairwise similarity estimates C₀₂ = 0.52, 95% CI = 0.38–0.66). The quadrats under closed canopy had high diversity when weighting rare species (species richness), whereas quadrats with one or two gap opening records during the past 20 years had high diversity when weighting the abundance of species. Although canopy openings provided establishment conditions for saplings of some shade-intolerant species, due to the nature of small gap size, such habitats do not favor the dominance of shade-intolerant species. Even in a frequently disturbed forest, species composition and richness of recruited saplings were mainly contributed by shade-tolerant species. Although multiple canopy openings facilitated the establishment of shade-intolerant species, species diversity in the study forests is possibly mainly mediated by coexistence mechanisms of those shade-tolerant species rather than light-gap-related species strategies.