Subcanopy gaps in temperate and tropical forests

Abstract We present a model of gaps in the vertical structure of forest vegetation. The traditional model of a forest gap assumes the existence of a ‘hole’ in the uppermost canopy layer, often extending down to near the ground. The present model extends the concept to gaps at any level, including those in lower layers below an intact canopy or subcanopy. It assumes that gaps at any level represent spaces with unused resources, especially favourable for plant growth and survival. Evidence from temperate and tropical forests indicates that gaps in the subcanopy and understorey layers below intact canopies are common, and that plants have higher growth rates in them than in non-gap sites. We also extend this model to below-ground gaps in the root zone.     

德国南部落叶阔叶林下常见植物的光适应性研究：—以德国Kraichtal地区落叶阔叶林为例

通过在13个样地中测定林下相对照度和多个植物物种的盖度指标,建立了林下相对照度与林下植盖度变化的相关关系,对德国南部落阔叶林下12种植物的光适应性进行了分析。结合植物的生态因子指示值和种在群落中的恒有度等生态指标,探讨了林下植物群落地位的形成原因。结果表明：1）在散射光条件下进行森林群落中的光照分布特征测定,能够降低光斑造成的干扰,获得相对稳定的光分布;2）在被研究的12个物种中,有9种植物在林下的分布与光照条件相关显,其中7种植物的盖度与相对照度之间具有正相关关系;2种植物具有负相关关系。其科3种植物的盖度与相对照度之间无显的线性相关;3）与光指示值相比,盖度与照度之间关系的定量分析能够更清楚地反映林下植被的光适应性,揭示植物的光生态幅特征;4）光适应性直接影响着林下植物的种间关系,也影响着物种的群落地位和作用。光生态幅广的植物,在群落中具有较强的适应能力,因此能够在多种群落类型中出现,表现出较高的恒有度值。     

Study on the Adaptation of Certain Species to Light Condition Under the Deciduous Broad-leaved Forests in Southern Germany?aTaking the Forest Communities in Kra ichtal,Germany as an Example

This study was conducted at the Kraichtal, southern Germany. Through the measurements of the relative illuminance under canopy and the coverage of many undergrowing species in 13 plots of forest communities, the correlation between the relative illuminance and the coverage of the species was calculated. The adaptation of 12 undergrowing species to light condition was determined based on the above calculation. Combining the indicator values of the species to ecological factors like light, humidity, temperature, acidity and nitrogen content in the soil with the constancy values of species in communities, the ranking order of undergrowing species in the communities was also examined. The results show: 1) In determining the light distribution in a forest community, scattered light condition is preferable because the interference from the drifting of light spots under the tree canopy can be reduced, and thus relatively stable distribution of light radiation can be obtained. 2) The distribution of 9 species, out of 12 species studied, was linearly correlated with the light condition under the canopy. For 7 of the 9 species, the correlation between the coverage of species and the relative illuminance was positive, and negative for 2 of them. For the other 3 species under the canopy the coverage was not correlated with the relative illuminance. 3) Compared with the light indicator values of species, the quantitative analysis of the relationships between species coverage and relative illuminance is more effective to reflect the adaptation of the species to light conditions under canopy and to reveal the ecological range of illuminance level of the species. 4) The relative adaptability of different species to light conditions directly affects the inter-specific relationships of the undergrowing species and may determine their ranking order in a community. The species with euphotic ecological range of illuminance level were easier to adapt to different light conditions under tree canopy and thus were able to exist in most forest community types with high constancy values.     

三种乡土树种在二种林分改造模式下的生理生态比较

在丘陵荒山先锋树种马占相思林的林窗和均匀间伐50％(简称林冠层下)的2种林分改造模式下,研究了降真香、红椎和火力楠3种乡土树种生长初期的生理生态。结果表明,不管是在冬季或夏季,林窗中比林冠层下有更高的光合有效辐射(PAR)和相对湿度(RH),并且有较低的气温(Tair),林窗与林内最主要的差异是PAR的变化。生长在林窗里的植株比林冠层下的叶片小而厚,叶片单位面积干重增加,林窗里的植株叶片有较高的叶绿素含量。冬天,降真香和红椎在林窗里和林冠层下的净光合速率(Pn)日变化曲线都为单峰型,而火力楠在林窗下为双峰型,林冠层下为单峰型。3种树种在林窗里和林冠层下植株的蒸腾速率(Tr)有较相似的日进程,都为单峰型。2种林分改造模式下3种树种叶片气孔导度(GS)的日变化也较相似,与PAR的变化趋势相似,而与RH变化呈相反趋势。冬季。降真香和红椎在林窗的水分利用效率(WUE)比林冠层下稍高,而火力楠在林冠层下则比林窗高．但它们都无显著差异。夏天,3种树种在林窗和林冠层下的Pn与冬天有相似的结果。夏季红椎和降真香在林窗的Tr比林冠层下高,而火力楠在林冠层下的Tr比林窗高。Gs的变化趋势也与PAR相似,与RH变化趋势相反。夏季,降真香和火力楠的WUE在林窗比林冠层下高,但红椎却比林冠层下的低。3个树种中火力楠的WUE最高。3种树种在林窗和林冠层下的wUE都是冬季比夏季高。综合植物各项生理生态指标的分析结果可知,降真香和红椎较适宜种植在林窗里,而火力楠较能适应有一定郁闭度的林冠层下。     

格氏栲天然林林窗植物群落功能性状的变异

林窗是森林更新演替的重要环节, 揭示林窗环境下功能性状变异来源及其相对贡献, 有助于阐明植物对林窗环境的响应。该研究以中亚热带格氏栲(Castanopsis kawakamii)天然林为对象, 设置9个不同大小的林窗样地, 运用方差分解探讨林窗、物种和个体对叶性状变异的相对贡献, 采用线性回归分析不同大小林窗下群落性状变化及种间和种内性状变异的重要性。研究发现: (1)格氏栲天然林林窗植物比叶面积、叶干物质含量、叶厚和叶绿素含量由种间性状变异主导, 叶氮含量由种内性状变异主导, 叶磷含量受林窗大小影响最大。(2)群落叶磷含量与林窗大小具有显著正相关关系, 土壤温度和水解氮含量对群落叶磷含量具有显著正效应, 土壤有效磷含量具有显著负效应。(3)沿林冠开放度的群落叶磷含量变化主要由种内性状变异引起, 优势种扮演着重要角色。结果表明, 格氏栲天然林林窗环境下植物功能性状仍以种间性状变异为主(平均41%), 但沿林窗环境梯度的群落性状变化主要源自种内性状变异, 通过植物表型可塑性响应环境改变, 优势种作用明显。     

油樟幼苗对马尾松林窗面积的光合响应特征

为了解马尾松人工林窗对伴生树种的影响,为马尾松人工纯林的团块状混交提供科学依据,研究了10 m×10 m(T1)、15 m×15 m(T2)、20 m×20 m(T3)、25 m×25 m(T4)、30 m×30 m(T5)、35 m×35 m(T6)和40 m×40 m(T7)马尾松人工林窗中油樟(Cinnamomum longepaniculatum)幼苗叶片形态和光合生理特征的变化,探讨马尾松林窗斑块对混生树种生长的影响。结果表明:1)林窗面积低于20 m×20 m时,油樟幼苗叶片最大净光合速率显著低于旷地对照;2)叶片比叶重随着林窗面积的增大显著升高;林窗内油樟幼苗叶氮含量在小林窗中(10 m×10 m)显著低于旷地对照,但在大林窗中(如20 m×20 m)显著高于对照;林窗内幼苗叶磷含量则与旷地无显著性差异;3)叶氮在光合组分中的总分配系数随着林窗面积的增加而增大,其中叶氮在羧化组分中的分配系数升高尤为明显,而捕光组分的分配系数在林窗面积10 m×10 m—20 m×20 m范围内随林窗面积的增加而显著降低。可见,当马尾松林窗面积低于20 m×20 m时,林窗环境会显著影响油樟幼苗的光合能力,油樟幼苗可以通过调节比叶面积、叶氮含量以及叶氮在光合组分中的分配等形态、生理适应特征来适应林窗环境的变化。     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

Differential effects of anthropogenic edges and gaps on the reproduction of a forest‐dwelling plant: The role of plant reproductive effort and nectar robbing by bumblebees

Forest fragmentation produces sharp changes in the spatial configuration of remnant forest patches, which include the increasing influence of patch edges on the interior of forest patches (‘edge effects’). Human activities responsible for forest fragmentation tend also to change the internal characteristics of remnant patches, for example, through the creation of gaps by selective logging. While edges and gaps can be expected to cause comparable changes in the micro‐environmental conditions of the forest, their effects on forest‐dwelling species and their interactions are not necessarily comparable. This study compares the effect of forest edges and anthropogenic gaps on the reproductive success of a self‐incompatible epiphytic plant (Mitraria coccinea), mediated by changes in its relationship with mutualists (pollinators) and antagonists (flower larcenists). Mitraria coccinea's flowers are pollinated by hummingbirds and robbed by the bumblebee Bombus dahlbomii. Edges and gaps had comparable positive effects on flower production and fruit set; however, nectar robbing was up to sevenfold higher in patches with numerous gaps and resulted in lower reproductive success (fruit set and total fruit crop). Forest fragmentation studies should therefore avoid treating forest remnants as homogeneous units (i.e. focusing exclusively on their characteristics and connectivity), without taking into account the internal heterogeneity caused by anthropogenic pressures (e.g. gaps and edge effects).     

Seed predation patterns favor the regeneration of dominant species in forest gaps compared with the understory in an oak-pine mixed forest

Recent studies have demonstrated the higher likelihood of regeneration in forest gaps compared with the understory for the dominant species in pine-oak mixed forest. Here, we tested whether rodent seed predation or dispersal was beneficial for gap regeneration. We tracked the seed predation and dispersal of Quercus aliena var. acuteserrata and Pinus armandii using coded plastic tags in the forest understory close to gaps. Our results demonstrated that the proportions of initial buried seeds of both species were significantly more abundant in the forest understory compared with gaps. After seed caching, however, significantly lower proportions of the seeds of both species survived in the forest understory compared with gaps during the 30-day observation period. The final survival proportions of the seeds cached in the forest understory were lower than those cached in the gaps the next spring, which indicated that small rodents rarely retrieved scatter-hoarded seeds from forest gaps. Our findings suggest that rodent seed predation patterns contribute to the regeneration of the dominant species in gaps compared with the understory in a pine-oak mixed forest. In the study area, reforestation usually involves planting seedlings but direct sowing in forest gaps may be an alternative means of accelerating forest recovery and successional processes.     

格氏栲林林窗自然干扰规律

通过对格氏栲(Gastanopsis kawakamii)林林窗自然干扰的基本规律的研究,得到了描述林窗干扰状况的一些重要特征。结果表明：格氏栲林中扩展林窗(EG)所占面积比例为34．46％,而实际林窗(CG)所占的面积比例为17．81％,干扰频率每年分别为0．69％和0．36％,林窗干扰返回间隔期为281a;EG的大小变化为63～1257m^2之间,平均为244m^2,而CG的大小变化为22～707m^2之间,平均为126m^2;形成林窗的主要方式是干中折断,比例占36．22％,其次为枯立,比例占32．28形;大多数的林窗是由1～5株形成木形成,3株形成木形成的林窗最多;大多数林窗是在前50a形成的,其中10～40a之前形成的林窗最多;林窗分布格局是均匀分布型的。林窗形成木主要由格氏栲、大叶乌饭(Vaccinium mandarinorum)、山胡椒(Lindera glauca)、狗骨柴(Tricalysia dubi)组成。林窗形成木的地径主要集中在60cm以下,地径20～60cm的主林层乔木形成林窗的比例较大。     

Beetle assemblages in rainforest gaps along a subtropical to tropical latitudinal gradient

Successional processes in forest gaps created by tree falls are often considered a principal mechanism maintaining species diversity within forests. Although insects are important as mediators of forest recovery processes, there have been few observations of how they differ between forest gaps and non-gaps across latitudes. Here we tested the impacts of gap habitat on beetle assemblage composition at three locations spanning sub-tropical and tropical biomes (28°S, 16°S, 5°N: Lamington and Daintree in Australia, and Danum in Borneo). Beetles were collected by flight interception traps and sorted to family and feeding guilds. Despite clear differences in environmental conditions between gaps and non-gaps at all three location, we found weak and inconsistent differences in beetle family and feeding guild composition across a latitudinal gradient. PERMANOVA results showed no significant differences in beetle family and feeding guild composition between gaps and non-gaps. For beetle families, however, the interaction effect (between gap vs. non-gap treatment and location) was significant. Post-hoc analysis and NMDS ordinations showed significant and clear separation of family assemblages between gaps and non-gaps within the Danum samples but not within samples from either the Lamington or Daintree sites. Using IndVal, when all three locations were combined, Chrysomelidae and Tenebrionidae were found to be indicators for gaps and Scarabaeinae and Leiodidae for non-gaps. Analysed separately, gap indicators were only Chrysomelidae at Daintree, whereas non-gap indicators were Scarabaeinae at Lamington, Scarabaeinae, all other Scarabaeidae, Anthicidae, Scydmaeninae, and Hybosoridae at Danum. We suggest the scale of changes in insect composition and richness with the creation of a natural light gap in forests contrast with those resulting from anthropogenic disturbance such as logging and clearing of forests and the greater adaptability of forests to small-scale changes rather than large-scale changes.     

洪水漫溢对林窗微环境时空差异的影响——以塔里木河中游荒漠河岸林为例

林窗是森林生态系统中普遍存在的干扰形式。探究洪水漫溢对林窗内部微环境时空异质性的影响,对揭示荒漠河岸林物种多样性变化及其更新特征具有重要意义。在塔里木河中游非洪水和洪水漫溢区各选取一个大小相似的林窗样地,并使用便携式气象监测仪对林窗内不同方位的空气温湿度进行监测,对比分析不同水淹条件下林窗内微环境的时空分布差异,为深入探索荒漠河岸林植被更新机制奠定基础。结果表明：（1）非洪水漫溢区和洪水漫溢区林窗内空气温度均呈现先上升后下降的变化趋势,空气湿度均呈现先下降后上升的变化趋势,且同一个样地中温度与湿度之间存在明显的负相关关系,而洪水漫溢改变了森林微环境,使林窗内湿度升高,温度下降。（2）非洪水漫溢区和洪水漫溢区林窗内不同方位温度分布差异较小,洪水漫溢对温度变化影响不大;两个样地林窗内湿度变化过程较为复杂,差异明显,洪水漫溢区湿度变化梯度更为密集。（3）非洪水漫溢区和洪水漫溢区林窗内温度差界限明显,西南方向温度差较小,中心位置温度差最大,洪水漫溢并未改变不同方位温度变化趋势;湿度差以中心偏北方向较大,其中非洪水漫溢区湿度差在西北方向较小,而洪水漫溢区湿度差最小值出现在西南方向。研究结果表示荒漠河岸林林窗微环境时空差异具有干旱区独特性,同时阐明了林窗微环境对洪水漫溢的响应,为深入研究林窗干扰对荒漠河岸林更新与演替提供了科学依据。     

Spatial and temporal variability of canopy structure in a tropical moist forest

Tree fall gaps are widely considered to play a prominent role in the maintenance of species diversity, while the spatiotemporal variability of canopy structure within closed forest stands is largely ignored. In this study we examined the vertical and horizontal components of canopy structure and its seasonal variability in a tropical wet semideciduous rainforest in Panama. Leaf area indices (LAI) were derived from measurements of diffuse radiation and empirically-based leaf angle distribution by mathematical inversion of a light interception model. Vertical distribution of LAI was non-homogeneous with 50% of the leaf area being concentrated in the uppermost 5 m of the canopy. In the wet season, when foliage is most abundant, the horizontal distribution of LAI in a 2100 m² plot ranged widely from 3 to 8, with a mean of 5.41. Changes in mean LAI between wet and dry seasons were small but highly significant. While ca 40% of the area was not affected by local changes in LAI, sizeable small scale changes in LAI did occur between wet and dry season in some locations. Local changes in LAI ranged from –2.3 to 2.4. These changes resulted in a 50% or more increase in light reaching the forest floor at 29% of the measuring locations, and a doubling or more at 13% of the location. Our results imply that structural heterogeneity by simple tree fall gaps do not adequately describe the dynamics of forest canopies.     

A role-type model (rope) and its application in assessing climate change impacts on forest landscapes

Gap-phase replacement is a general phenomenon found in forest ecosystems, worldwide. Different tree species can be expected to produce different sizes of gaps when they die. Species also vary in their regeneration success in gaps of different sizes. In this paper, the gap-phase interactions among tree species in a forest stand are simulated by a role-type stand model called ROPE. By incorporation of environmental effects on tree height, ROPE can simulate forest composition and stand leaf area under different climate conditions. The model was developed for forest ecosystems in northeastern China and was used to simulate the forest landscape structures under current climate conditions and under four climate change scenarios for greenhouse gas related warming. These scenarios were obtained from general circulation models developed by different atmospheric research centers. Korean pinebroadleaf mixed forest and larch forest are the major stand types in the study area under present conditions. Under the four climate change scenarios, Korean pine-broadleaf mixed forest would be expected to occur only on the higher parts of large mountains. Larch forest only would be found north of the study area. Broadleaf forest would become the dominant vegetation over the study area. Use of the Kappa statistic to test for similarity in spatial maps, indicates that each climate change scenario would result in a significant change of forest distributions.Supported by The United States National Science Foundation Grant BSR-8702333 to University of Virginia.     

Treefall gap characteristics and regeneration in the laurel forest of Tenerife

Abstract. We conducted a study in the laurel forest of Tenerife (Canary Islands, Spain) to describe the characteristics of natural gaps and to assess the role of treefall gaps in forest dynamics. Very little is left of the natural laurel forest with i.a. Laurus azorica, Ilex canariensis and Prunus lusitanica. We looked for treefall gaps in 80 randomly located 2500 m² plots. These plots represented ca. 1% of the remaining and protected laurel forest of Tenerife. We recorded the characteristics of the species causing the gaps, gap architecture and gap age in all observed gaps larger than 10 m². We inventoried the regeneration in each gap and in a neighbouring control plot with the same topography. Large gaps (>75 m²) were not common in the laurel forest. The absence of large gaps could be due to the physiognomy of the vegetation, the mild weather or the rarity of disturbances. Instead of forming gaps, many trees decompose in place and branches from neighbouring trees and suckers from the decomposed trees occupy the free space. Also, the high rate of asexual regeneration could contribute to the fast closing of the gap. The number of gaps created by Prunus lusitanica was higher than expected (based on canopy composition) while Ilex canariensis and Laurus azorica created fewer gaps. In this evergreen forest, differences between gap and non-gap conditions are not as distinct as in other forest types. Only 0.4% of the canopy is in the gap phase (0.6% including gaps smaller than 10m²). No differences were found in patterns of regeneration between gap and non-gap phases in the forest. Gaps do not explain the persistence of pioneer species in the laurel forest.     

格氏栲天然林林窗和林下种子散布及幼苗更新研究

以格氏栲(Castanopsis kawakamii)天然林为研究对象,探讨了林窗和林下格氏栲种子雨、种子库的分布特征及幼苗更新状况。结果表明:林窗种子雨总量和完好种子密度高于林下,未成熟种子比例低于林下;林窗和林下种子雨高峰期掉落数量分别占种子雨总量的77.13%和74.5%;林窗种子库储量低于林下,种子库中以全食或捡拾种子比例最高,其中种子库储量垂直分布表现为枯落物层(约占2/3)>腐殖质层(0～5 cm)(约占1/3)>心土层(5～10 cm)(小于1%),以格氏栲种子占绝对优势;格氏栲从种子到幼苗的转化率低,林窗中格氏栲完好种子密度与幼苗密度均高于林下。林窗微生境提高了种子散布过程中格氏栲成熟种子的密度和比例,有利于促进格氏栲的幼苗更新,表明林窗在格氏栲种群恢复过程中扮演着重要角色。     

Modelling Direct Radiation and Canopy Gap Regimes in Tropical Forests

Spatial and temporal variation in the below‐canopy light environment of tropical forests is not well known and its measurement is technically challenging. Distributions of gap and understory areas in forests are likewise little known because of the resource requirements of forest structural censuses and a lack of consensus over how gaps should be defined. A basic model of forest structure, based on tree allometries from the 50 ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and a solar positioning algorithm were used to predict spatial and temporal variation in the distribution of direct light at the forest floor. Predicted duration of direct sunlight was then compared with the distribution of gap and understory areas, delimited according to four standard gap definitions, giving predictions for the correspondence between direct light regimes and forest structure. At least 36 percent of the areas of gaps of all sizes was predicted to receive < 1 h of direct sunlight per day, and the understory to receive direct sunlight for ≥ 1 h per day in up to 15 percent of its area, even when not in proximity to gaps. The predicted distribution of light changed over the course of the year with the greatest spread of light throughout the forest floor coinciding with the months when maximum daily solar elevation peaked. These predictions suggest a partial decoupling of light regimes from canopy structure, with implications for gap definitions, patch models of forest development and current understanding of tree seedling recruitment patterns.     

林窗尺度对侧柏人工林土壤微生物量和功能多样性的影响

微生物活性是影响土壤碳循环等地下生态系统过程的重要因素。以徐州市侧柏(Platycladus orientalis(L.)Franco)人工林为研究对象,以未受干扰的侧柏林为对照(CK),设置半径分别为4、8、12m的3种尺度近圆形林窗,从林窗边缘(D1)到距林缘4m(D2)及8m处林下(D3)水平梯度上分析土壤微生物量碳(MBC)、氮(MBN)和代谢功能多样性的变化。结果表明:1)与CK相比,林窗样地土壤MBC总体降低,MBN含量显著下降(P0.05),MBC/MBN显著上升(P0.05)。在3种尺度林窗中,MBC在大林窗偏小,MBN在小林窗偏小;MBC/MBN总体上随林窗尺度增大而减小。2)与CK相比,大中林窗降低了土壤微生物代谢活性(AWCD),小林窗变化不大。从D1到D3点,小林窗的AWCD先降后升,中林窗呈上升趋势,大林窗则相反。而且林窗降低了土壤微生物对各类碳源的利用,主要利用聚合物类和氨基酸类碳源,中林窗样点对碳水化合物和氨基酸类小分子碳源的利用最低。3)林窗总体提高了土壤微生物功能多样性,其中多样性(H')、丰富度(S)和均匀度(E)3个指数在各点之间均无显著差异,小林窗和CK的优势度指数(D_s)显著大于(P0.05)大林窗。侧柏林人工林窗对土壤微生物量和功能多样性的影响有着明显的尺度和位置梯度效应,林窗有望促进侧柏林土壤碳固持和大分子物质降解,提高其应对全球气候变化的能力,综合而言,中尺度林窗对侧柏林生态功能的发挥较为有利。     

基于无人机影像探讨格氏栲天然林林窗数量分布及其影响因素

探讨格氏栲(Castanopsis kawakamii)天然林林窗数量特征及其空间分布对预测森林种群动态变化及演替具有重要意义。该文采用无人机航拍获取格氏栲天然林正射影像图并结合野外调查,通过提取林窗特征参数和计算植被覆盖率来探讨林窗空间分布及其影响因素。结果表明:(1)保护区内格氏栲天然林植被覆盖率为75.53%,部分地区出现一定面积裸露土地。(2)研究区林窗空隙率为2.40%,密度为6.50ind.·hm-2,平均林窗面积为36.86 m2。(3)研究区林窗数量随林窗面积增加呈负指数分布,以微、小和中型林窗为主,面积100 m2以上的林窗数量较少。(4)低海拔林冠层覆盖度好,中海拔地区林窗个体数和平均林窗面积较大,高海拔地区林窗密度和空隙率相对较大。林窗主要分布在缓坡和斜坡上,其平均面积、密度和林窗空隙率也明显较高。西和南方位林窗数量较多,北、西北和东南方向林窗密度和空隙率相对较大。格氏栲天然林植被覆盖率较高,以微、小和中型林窗为主,地形因子通过改变林窗面积、林窗密度和林窗空隙率特征驱动了格氏栲天然林林窗数量与空间分布格局。     

Spatial patterning in the canopy gap regime of a subalpine Abies-Picea forest in the northeastern United States

Abstract. Spatial patterning in the disturbance regime of a forest affects the vegetation dynamics. Therefore, the distribution of canopy gaps was examined in detail for a subalpine Abies-Picea forest in the northeastern United States. Gaps were not randomly distributed. The fraction of forest area in gaps and the abundance of gaps varied significantly with topographic position, elevation, and slope percent. On average, 15 % of the forest was influenced by gaps, but the gap fraction was greater near ridges (23 %) and near streams (27 %) than on the backslope (13 %). Also, gaps were larger and more abundant near streams and ridges. Gap fraction varied with elevation as well: more of the forest was disturbed at lower and higher elevations than at mid-elevations. Significantly more of the forest on steep slopes (≥ 30 %) was under gaps. As a result of this patterning, some parts of the Abies-Picea forest were predictably more disturbed than others. A remaining question is whether this patterning is sufficient to influence the regeneration environment and thus forest composition.