A modular approach to biomass allocation in an invasive annual (Microstegium vimineum; Poaceae)

The invasive grass Microstegium vimineum grows in low light beneath the canopy of eastern forests in North America by reiteration of modules (phytomers) along a tiller. Basal phytomers are vegetative; terminal phytomers produce a raceme of chasmogamous (CH) spikelets plus an axillary raceme of cleistogamous (CL) spikelets. Additional subterminal phytomers with CL racemes mature basipetally. Allocation to culms, leaves, and CH and CL within phytomers was examined in relation to light conditions for a population in New Jersey, USA. Plants were reared in a greenhouse from seed families of parents in deep shade (2-8% full sun) or sunny, edge habitats. Primary tillers were subdivided into phytomers, dried, and weighed. Tillers from field habitats were similarly treated. For vegetative and subterminal phytomers, allocation to leaves and CH was greatest for the shady habitat. CL allocation decreased from terminal to reproductively immature subterminal phytomers. CH and CL mass was positively correlated with leaf mass, suggesting that reproduction is determined by available photosynthate. CH mass showed a genetic correlation with leaf mass. Developmental plasticity in modular allocation allows Microstegium to maximize fitness when conditions are favorable (e.g., high light along forest edges) by continual maturation of CL caryopses on axillary racemes.     

Growth trajectories and size-dependent reproduction in the highly invasive grass <Emphasis Type="Italic">Microstegium vimineum</Emphasis>

To better understand the ecological and evolutionary processes that underlie the ability of invasive plants to colonize and spread in a region, basic data on quantitative genetic variation in growth and reproduction of plants from contrasting microsites are desirable. The annual grass Microstegium vimineum is an invasive colonizer of the herbaceous layer in forests of the eastern United States. This greenhouse study utilized seeds collected from maternal plants in a shady forest or sunny edge microsite in central New Jersey, USA, to examine variation in growth trajectories and size-dependent reproduction. Questions addressed were (1) do growth trajectories vary significantly among families and between microsites? and (2) does reproductive mass scale with vegetative size? The trajectory for shoot dry mass over time was significantly related to microsite, due apparently to greater growth over the final 8 weeks of plants from the interior compared to those from the edge. It is suggested that a growth increase late in the season enables plants to maximize reproduction when light conditions improve following canopy leaf senescence. Number of tillers exhibited significant variation among families within microsites, but growth trajectories did not. Positive phenotypic and genetic correlations were detected between reproductive and vegetative mass per tiller; thus selection may favor large tillers to increase seed output, a critical life history trait for an invasive weed.     

The putative niche requirements and landscape dynamics of Microstegium vimineum: an invasive Asian grass

The theoretical foundations of population and community ecology stress the importance of identifying crucial niche requirements and life history stages of invasive species and, in doing so, give insight into research and management. We focus on Microstegium vimineum, an invasive grass which is causing marked changes in the structure and function of US forests. We describe M. vimineum’s life history and habitat characteristics, infer its niche requirements and synthesize this information in the context of population dynamics and management. Based on the results synthesized here, M. vimineum’s crucial niche requirements appear to be light (reproductive output), soil moisture (reproductive output, seedling recruitment) and aboveground coverage by leaf-litter and competing species (seedling recruitment and survival). These data suggest a source-sink dynamic might allow M. vimineum to disperse and thrive along sunny, and sometimes wet, edge habitats and, in turn, these populations might act as source populations for adjacent shady forest habitats. By evaluating M. vimineum in the context of its stage-specific requirements, we highlight potential weaknesses in its life history that provide strategies for effective management.     

VARIATION IN THE DEGREE OF CLEISTOGAMY WITHIN AND AMONG SPECIES OF THE GRASS DANTHONIA

North American species of the grass Danthonia bear distinct chasmogamous and cleistogamous flowers. Four taxa occurring in North Carolina (D. compressa, D. epilis, D. sericea, and D. spicata) were examined for variation in the degree of cleistogamy. The flowering culms bear a single terminal panicle consisting of 4 to 12 spikelets of chasmogamous flowers. In each leaf axil, at the nodes of the flowering culm, is a single spikelet of cleistogamous flowers completely surrounded by the leaf sheath. The percentage of cleistogamous flowers produced on any culm depends on the number of chasmogamous and cleistogamous spikelets and the number of flowers in each type of spikelet. All four characters vary among the taxa examined. Danthonia compressa produces, on the average, 50% cleistogamous flowers; D. spicata, 25% cleistogamous flowers; D. sericea and D. epilis, 5% cleistogamous flowers. The degree of cleistogamy in D. spicata is associated with certain habitat features. Populations from mountain sites, disturbed sites, and non-woodland sites produced higher percentages of cleistogamous flowers than did populations from piedmont, undisturbed, and woodland sites, respectively. Grazing may favor increased cleistogamy because cleistogamous flowers are produced lower on the plant. In North Carolina, the most frequently grazed Danthonia taxa also produce the highest percentage of cleistogamous flowers.     

Changing light conditions in pine rockland habitats affect the intensity and outcome of ant–plant interactions

Extrafloral nectar (EFN) mediates food‐for‐protection mutualisms between plants and ants. Such mutualisms exist within a complex web of biotic interactions, and in a framework provided by the abiotic environment. Both biotic and abiotic factors, therefore, affect the outcome of ant–plant interactions. We conducted an experiment to determine the effects of ant activity, and light intensity, on herbivory rates, growth, and reproductive fitness in Senna mexicana var. chapmanii, a perennial legume native to pine rockland habitats of south Florida. Forty plants were divided among four treatments in a factorial experimental design with two independent variables: ant activity and light intensity. Plants were divided equally between sunny and shady habitats, and ants were excluded from half of the plants in each habitat type. The presence of ants significantly reduced herbivory rates in S. chapmanii. In shaded habitats, the presence of ants had no effect on plant reproductive fitness, however, in sunny habitats plants with ants produced significantly more seeds over the duration of the 1‐yr study. Ants represent an important biotic defense against herbivores in S. chapmanii; however, their effects on plant fitness are dependent on light conditions. Pine rockland habitats in south Florida have been widely destroyed or mismanaged. In fragments that remain, suppression of fire has led to increased canopy closure and shading of the understory. These changes will likely negatively impact plants that rely on ants for defense. We highlight the importance of conservation efforts to preserve the pine rocklands and the fire regimes on which they rely.     

Flowering Phenology and Gender Variation in Pennisetum typhoides

Because of the modular structure of pearl millet (an annual grass crop, Poaceae), different tillers of a plant share the same genotype but are subjected to different environmental conditions during their maturation. This allows investigation of the effects of tiller flowering phenology on allocation to resource-producing photosynthetic biomass, sexual functions, and thus tiller gender. All tillers of plants of two families collected from individual maternal plants (represented by 33 and 31 plants each) were analyzed. In both families, allocation to aboveground vegetative biomass decreased as flowering was delayed. On average, late-flowering tillers were 65% smaller than the first ones to flower. The proportion of biomass allocated to reproduction significantly increased with the flowering rank of the tillers, suggesting that translocations of assimilates occurred between early- and late-flowering tillers. In both families, late-flowering tillers produced significantly fewer pollen grains per stamen than early-flowering ones, and female reproductive allocation (expressed as seed mass per tiller) was also affected by flowering phenology. Tillers became increasingly female as flowering phenology progressed. This gender variation is possibly adaptive because pollination efficiency is maximized by plant height. Natural selection may favor a shift toward femaleness to maximize reproductive fitness in small, late-developing tillers.     

Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

TILLERING AND MORTALITY OF THE SALT MARSH CORDGRASS,SPARTINA ALTERNIFLORA

Recruitment and mortality of Spartina alterniflora tillers and culms were estimated from measurements of live standing crops at Wallops Island, Virginia. Stands of tall and medium form grass produced from 452 to 1,227 tillers m⁻² yr⁻¹. Those tillered in spring lengthened into culms their first year, while those tillered in late summer and fall may have overwintered and lengthened into culms their second year. Biennial growth was most prevalent in tall form Spartina. Fifty-six to sixty-one percent of all tillers died before lengthening into culms. Of the remainder, 9 to 56% died as culms before the time of flowering. Sixty-three percent of the surviving culms of tall form Spartina flowered, compared to only 14 to 23% in medium form. Mortality of tillers and culms removed 16 to 35% of the annual production of Spartina biomass from live standing crops before the time of flowering.     

Neighbour influence on the tiller demography of two perennial pampa grasses

Abstract. The aim of this study was to evaluate the possibility to use neighbour species composition to explain demographic variations in seasonal tillers in perennial grasses. Two warm-season species of the Argentina flooded pampa, Paspalum dilatatum and Sporobolus indicus, were used as models. Tiller structure and dynamics of target plants and the identity of their nearest neighbours were analyzed within a natural grassland community for an entire annual growth cycle. Canonical Correlation Analysis of tillering behaviour of target plants and neighbour species composition showed significant relationships for summer neighbourhood data. Community components affecting tiller demography were identified, and their spatial variation was described as different gradients of neighbourhood composition (NCG). NCG varied between target species, determining different spatial variation in neighbouring canopy dynamics and tiller dynamics. The tiller structure and dynamics of P. dilatatum plants were significantly correlated with the seasonal pattern of their neighbouring canopy density (NCD). Plants placed on the NCD-fluctuating extremes along NCG showed an unstable modular structure and short-lived tillers, as compared to those placed in the centre of the gradient. Relative density of the neighbouring canopy showed little variation along the NCG of S. indicus plants, which presented a tiller structure more stable and uniform than P. dilatatum plants did. Identification of NCG may be a promising approach to understanding changes in tiller dynamics of target species in relation to spatial and temporal changes in community structure.     

色季拉山急尖长苞冷杉林不同坡向土壤养分及肥力研究

以色季拉山不同坡向急尖长苞冷杉(Abies georgei var.smithii)天然林为研究对象,采用单因素方差、化学计量比及内梅罗指数等方法对不同坡向0—100 cm土壤的pH和7种养分含量及肥力状况进行研究,以期为色季拉山急尖长苞冷杉的经营管理提供理论依据。结果表明:(1)色季拉山急尖长苞冷杉林土壤pH7,为酸性。0—100 cm土层中除土壤全钾含量外,其他各养分含量及变异系数的均值均表现为:阳坡阴坡,且均属于中等变异程度。除全磷、全钾含量较低外,其他指标均处于"中等"及以上水平。(2)土壤有机质、全氮、碱解氮随土层的增加而降低,总体上表层与深层差异显著(P0.05),其他土层之间差异不显著(P0.05);其他养分随土层变化规律不一致,各土层之间差异不显著(P0.05)。(3)阳坡各土壤化学计量比总体上表现为随土壤深度增加而降低,阴坡呈波动性变化,除C∶N外,阳坡C∶P、C∶K、N∶P、N∶K、P∶K均小于阴坡。(4)阴坡的土壤肥力指数(1.44)大于阳坡(1.32),不同坡向土壤肥力均属于"中等"水平。坡向通过影响多种因素使土壤理化性质具有一定差异性,阴坡的土壤养分含量及肥力状况优于阳坡,全钾是第一限制因子,全磷是第二限制因子。     

桂林岩溶石山植物群落植物功能性状对不同坡向环境因子的响应

该研究以桂林岩溶石山不同坡向的植物群落为对象,探讨植物功能性状(比叶面积、叶绿素含量、木材密度和环境因子随坡向的变化规律。结果表明:比叶面积和叶绿素含量为阴坡>半阴坡>阳坡,阴坡和半阴坡分别与阳坡差异性显著;木材密度为阴坡<半阴坡<阳坡,阴坡、半阴坡分别与阳坡差异性显著。土壤有机质含量表现为阴坡大于阳坡,且阴坡与阳坡差异性显著;土壤全磷、土壤有效磷均表现为阳坡含量最高,且阴坡与阳坡差异性显著;土壤有效钾、土壤全钾则分别为阴坡、半阴坡含量最高,土壤全钾含量在各坡向上差异均显著,而土壤有效钾则为阴坡、半阴坡与阳坡均差异性显著。回归分析表明群落水平比叶面积在阴坡和半阴坡上与土壤有机质含量呈显著负相关;群落水平叶绿素含量与土壤全磷含量和土壤有效钾含量在阴坡上呈显著正相关;在不同坡向梯度上群落水平木材密度与环境因子无相关性。群落水平上植物功能性状随坡向的变化规律,反映了岩溶石山植物群落构建过程中环境对功能性状的筛选效应。     

Demographic variability in tiller populations of two perennial pampa grasses

Abstract. The general objectives of this study were: (1) to investigate the importance of internal influences in regulating the tiller dynamics in natural populations of the warm-season perennial grasses Paspalum dilatatum and Sporobolus indicus, coexisting in Argentine flooded pampa, in as much as they act independently of the underlying external environment, and (2) to evaluate the extent to which interactions between internal and external factors affect the variation in tiller dynamics within such populations. Within-population variation in seasonal development of plants and tillers with different neighbour composition was studied for an annual growth cycle. Tiller survival and tillering were significantly influenced by tiller size. Tiller age influenced tiller fate, as suggested by the additive effects of age and size of tillers. These relationships varied with season and with species. Size and age of tillers showed additive effects with their neighbouring species on the tiller fate of P. dilatatum, but the effects of age and size of S. indicus changed according their neighbourhood. Tiller survival of S. indicus during the early growth season was more size-dependent when the cold-season species Poa lanigera, was the principal neighbour. Flowering and tillering probabilities were positively related through their common positive responses to tiller size. Tiller survival and recruitment between different seasons were strongly related. Independently of neighbour composition, tiller survival was generally inversely related to tiller recruitment in previous seasons. Therefore, significant density-dependent mortality of tillers was found for both species during the early summer when tiller density was expressed by basal area units.     

Per capita community-level effects of an invasive grass, Microstegium vimineum, on vegetation in mesic forests in northern Mississippi (USA)

Quantifying per capita impacts of invasive species on resident communities requires integrating regression analyses with experiments under natural conditions. Using multivariate and univariate approaches, I regressed the abundance of 105 resident species of groundcover plants and tree seedlings against the abundance and height of an invasive grass, Microstegium vimineum, within 117 plots in four mesic floodplain forests in Mississippi (USA). Microstegium vimineum was most productive (i.e., tallest and most abundant) in canopy gaps in floodplains, and a significant amount of variation in resident species composition was directly explained by canopy gaps and stand age. The relatively small (but statistically significant) percentage of variation in resident species composition (1.8%) explained by M. vimineum in the multivariate analysis was attributable to significant relationships with a few common species. Most of these were negative relationships with shady mesic forest indicators. Most positive relationships were with infrequent disturbance indicators and with species with growth phenologies that differed from that of M. vimineum. Results of field competition experiments with the three most common species to show significant relationships with M. vimineum revealed asymmetric competitive effects of M. vimineum on Chasmanthium laxum and positive responses of Quercus alba seedlings and Leersia virginica adults to the removal of M. vimineum in one growing season. Results of this study suggest that negative per capita community-level effects of M. vimineum are likely to be greater in shady forests than in open floodplain forests due to the relative paucity of vulnerable species in the latter.     

Tillering in grain sorghum over a wide range of population densities: identification of a common hierarchy for tiller emergence,leaf area development and fertility

Most studies of tiller development have not related the physiological and morphological features of each culm to its subsequent fertility. This introduces problems when trying to account for the effects of tillering on yield in crop models. The objective of this study was to detect the most likely early determinants of tiller fertility in sorghum by identifying hierarchies for emergence, fertility and grain number of tillers over a wide range of assimilate availabilities. Emergence, phenology, leaf area development and dry weight partitioning were quantified weekly for individual tillers and main culms of tillering and uniculm plants grown at one of four densities, from two to 16 plants m(-2). For a given plant in any given density, the same tiller hierarchy applied for emergence of tillers, fertility of the emerged tillers and their subsequent grain number. These results were observed over a range of tiller fertility rates (from 7 to 91%), fertile tiller number per plant at maturity (from 0.2 to 4.7), and tiller contribution to grain yield (from 5 to 78 %). Tiller emergence was most probably related to assimilate supply and light quality. Development, fertility and contribution to yield of a specific tiller were highly dependent on growing conditions at the time of tiller emergence, particularly via early leaf area development of the tiller, which affected its subsequent leaf area accumulation. Assimilate availability in the main culm at the time of tiller emergence was the most likely early determinant of subsequent tiller fertility in this study.     

Tiller demography in seashore populations of Agrostis stolonifera,Festuca rubra and Poa irrigata

Abstract. The dynamics of tillers in natural populations of three cohabiting perennial grass species, Agrostis stolonifera, Festuca rubra and Poa irrigata (= Poa pratensis ssp. irrigata) were studied for five years in a Baltic seashore meadow. The process of tiller population maintenance was very dynamic. Both birth and death rates of tillers were high, particularly in A stolonifera, and the turnover rate of the populations was high. Recruitment was mainly by vegetative tillers, produced continuously throughout the growing season. The proportion of flowering tillers was low, but varied between years. Considerable year-to-year variation was also found in birth and death rates. Despite this between-year variation and the differences found between species in flowering frequency, pattern of survivorship and tiller longevity, population sizes of the species remained relatively constant.     

Tillering in grain sorghum over a wiide range of population densities: modelling dynamics of tiller fertility

The prediction of tillering is poor or absent in existing sorghum crop models even though fertile tillers contribute significantly to grain yield. The objective of this study was to identify general quantitative relationships underpinning tiller dynamics of sorghum for a broad range of assimilate availabilities. Emergence, phenology, leaf area development and fertility of individual main culms and tillers were quantified weekly in plants grown at one of four plant densities ranging from two to 16 plants m(-2). On any given day, a tiller was considered potentially fertile (a posteriori) if its number of leaves continued to increase thereafter. The dynamics of potentially fertile tiller number per plant varied greatly with plant density, but could generally be described by three determinants, stable across plant densities: tiller emergence rate aligned with leaf ligule appearance rate; cessation of tiller emergence occurred at a stable leaf area index; and rate of decrease in potentially fertile tillers was linearly related to the ratio of realized to potential leaf area growth. Realized leaf area growth is the measured increase in leaf area, whereas potential leaf area growth is the estimated increase in leaf area if all potentially fertile tillers were to continue to develop. Procedures to predict this ratio, by estimating realized leaf area per plant from intercepted radiation and potential leaf area per plant from the number and type of developing axes, are presented. While it is suitable for modelling tiller dynamics in grain sorghum, this general framework needs to be validated by testing it in different environments and for other cultivars.     

喀斯特退化天坑阴坡阳坡壳斗科植物的功能性状特征

喀斯特退化天坑的负地形生境中阴坡和阳坡的局域环境具有显著差异,导致植物群落类型差异明显。本研究以云南沾益退化天坑深陷塘为例,探究退化天坑阴坡与阳坡壳斗科植物功能性状特征,有助于揭示天坑的物种多样性保护库价值。结果表明: 阴坡土壤营养物质含量高于阳坡,阴坡壳斗科植物的叶面积、比叶面积和叶干重显著大于阳坡,叶厚度显著小于阳坡,叶干物质含量小于阳坡;阴坡壳斗科植物功能性状的主要环境影响因子是土壤全钾和土壤含水量,全磷是阳坡的主要环境影响因子。阴坡的叶功能性状变异程度小于阳坡,主要是以改变叶干重增加植物体内光合速率和碳积累能力的方式适应阴坡生境;阳坡通过保持较小比叶面积,增加叶面积变异程度来获得更多的资源。退化天坑深陷塘阴坡植物群落演替速度明显快于阳坡,壳斗科植物优势在阴坡逐渐减小,在阳坡仍是优势建群种。     

Limits to local spatial spread in a highly invasive annual grass (Microstegium vimineum)

The potential roles of seed and microsite limitation in local spatial spread of the invasive grass Microstegium vimineum were experimentally investigated in a woodland and open lawn in central New Jersey, USA. Plots (30 × 30 cm) in three sites previously unoccupied by M. vimineum for at least 8 years (woodland interior, woodland edge, and open lawn) were sown with ~262 M. vimineum seeds in early spring 2008. Seedling emergence, density, summer growth and autumn reproduction were compared to plots in a nearby control population where natural recruitment occurred. Seedling emergence was greatest in the open lawn (54% of seeds sown) where plants showed the greatest growth and reproduction due to high light availability. Seedling emergence was lowest in the woodland interior (24%) and edge sites (9%), and growth and reproduction were greatly reduced there (relative to the control). Plots in the open lawn supported a consistently high density of M. vimineum (>1,000 plants per m²) through the growing period (April to October). The number of seeds in both cleistogamous and chasmogamous spikelets was correlated with shoot dry mass and thus, total seed production was greatest in the sunny open lawn where plants were largest, despite high density there. Across all sites, plants in plots at the highest densities produced the most seeds. Total seed production correlated with levels of light, but not soil moisture. Both seed availability and microsite limitation may reduce the probability of establishment of new M. vimineum populations into previously unoccupied sites. Intraspecific density does not negatively affect survival or reproduction. Light and soil moisture can be limiting abiotic factors in some areas, but poor, natural seed dispersal limits the distribution of this invasive species on a local scale.     

Evolutionary significance of genotypic variation in developmental reaction norms for a perennial grass under competitive stress

The developmental reaction norm (DRN) represents the set of ontogenetic trajectories that can be produced by a genotype exposed to different environmental conditions. Genetic variation in the DRN for growth traits and in the patterns of biomass allocation is critical to phenotypic evolution in heterogeneous environments. The DRN and patterns of biomass allocation were investigated in 11 clones of the caespitose, corm-forming, perennial grass Phleum pratense in relation to competitive stress imparted by Lolium perenne in a 16 week glasshouse experiment. A separate experiment assessed the ability of basal buds flanking a corm to sprout and the relationship of corm mass to sprout mass for the same clones. Corm fresh mass varied among clones and was significantly correlated with the dry mass of the tillers that sprouted from basal buds. In the competition experiment, clones in competitive environments varied significantly from those in non-competetive environments in terms of their DRNs for number of tillers and shoot dry mass. Thus, selection of DRNs would favour different genotypes in the two environments and at different times. Significant negative genetic correlations were detected for tiller number and mean tiller mass in the noncompetitive, but not the competitive, environment. Biomass allocation to stem bases was significantly greater for clones under competitive stress. Allocation to storage tissues such as corms may be adaptive if it enhances persistence in the competitive field environments typically occupied by caespitose grasses. Root and shoot allocation showed a significant clone by competition interaction. For P. pratense, genotypic variation in growth trajectories plays an important role in determining variation in individual performance, a condition necessary for the continued evolution of the DRN.     

地形对典型阔叶红松林叶面积指数分布的影响

基于小兴安岭丰林国家级自然保护区阔叶红松林30 hm²监测样地,利用叶面积指数(LAI)与胸高断面积间的经验模型计算样地内主要树种及所有树种的总LAI,研究海拔、坡度、坡向和曲率4种地形因子对LAI分布的影响,以及对LAI分布的综合作用. 结果表明: 4种地形因子对红松、冷杉、紫椴、裂叶榆和五角槭5种主要树种LAI的分布均存在显著影响,而坡向对枫桦LAI的分布存在显著影响;4种地形因子对所有树种总LAI的分布具有显著影响. 将样地划分为山脊、阳坡、山谷阳面、阴坡和山谷阴面5种生境类型.生境类型对所有树种总LAI的分布存在显著影响,其中山脊LAI显著高于其他生境类型,均值为8.85;阳坡的LAI次之,均值为7.62;山谷阳面和阴坡的LAI不存在显著差异;而山谷阴面的LAI最小,均值为4.42.