Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Can isotropy vs. anisotropy in the spatial association of plant species reveal physical vs. biotic facilitation?

In dryland ecosystems and other harsh environments, a large part of the vegetation is often clustered, appearing as ‘islands’. If ‘independent’ species, usually colonizers, can be distinguished from species which are ‘dependent’ on the presence of the colonizing species for successful establishment and/or persistence, the type of spatial pattern of the association ‐ isotropic (spatially symmetric) or anisotropic (spatially asymmetric) ‐ can give information on the underlying environmental factors driving the process of association. Modified spatial pattern analysis based on Ripley's K‐function can be applied to bivariate clustered patterns by cardinal direction in order to detect possible anisotropy in the pattern of association. The method was applied to mapped distribution patterns of two types of semi‐arid shrubland in southeastern Spain. In shrubland of Retama sphaerocarpa, low shrubs of Artemisia barrelieri were significantly clustered under the canopy of the Retama shrubs in all four cardinal directions, suggesting an isotropic facilitation effect. In low shrubland dominated by Anthyllis cytisoides and Artemisia barrelieri, Anthyllis shrubs occurred more frequently than expected on the eastern side (and downslope) of an Artemisia shrub. The possible environmental factors driving the two association patterns are discussed and recommendations for further applications of the analytical method are given.     

Nutrient losses in runoff from grassland and shrubland habitats in Southern New Mexico: I. rainfall simulation experiments

Rainfall simulation experiments were performed in areas of semiarid grassland (Bouteloua eriopoda) and arid shrubland (Larrea tridentata) in the Chihuahuan desert of New Mexico. The objective was to compare the runoff of nitrogen (N) and phosphorus (P) from these habitats to assess whether losses of soil nutrients are associated with the invasion of grasslands by shrubs. Runoff losses from grass- and shrub-dominated plots were similar, and much less than from bare plots located in the shrubland. Weighted average concentrations of total dissolved N compounds in runoff were greatest in the grassland (1.72 mg/1) and lowest in bare plots in the shrubland (0.55 mg/1). More than half of the N transported in runoff was carried in dissolved organic compounds. In grassland and shrub plots, the total N loss was highly correlated to the total volume of discharge. We estimate that the total annual loss of N in runoff is 0.25 kg/ha/yr in grasslands and 0.43 kg/ha/yr in shrublands — consistent with the depletion of soil N during desertification of these habitats. Losses of P from both habitats were very small.     

Interaction between the cactus Neobuxbaumia tetetzo and the nurse shrub Mimosa luisana

Abstract. Pattern analysis and association analysis showed that recruitment of the giant columnar cactus Neobuxbaumia tetetzo in the semi-arid valley of Zapotitlán is largely limited to areas beneath the canopies of perennial shrubs, acting as nurse plants. Chi-square and Haberman tests revealed that young cacti were more frequently found beneath canopies of leguminous shrubs, especially Mimosa luisana, than were older cacti. Segregation analysis indicated a positive association (= negative segregation) of young cacti and M. luisana canopies, but older cacti were randomly distributed with respect to M. luisana. This, and the height class distributions of cacti associated withM luisana or not, suggested the replacement of M. luisana by N. tetetzo. Mimosa luisana plants in association withiV. tetetzo had greater amounts of dead basal area thanM luisana lacking associated N tetetzo. This suggested competition from JV. tetetzo promotes the replacement process. The roots of N tetetzo are ≤ 30 cm deep. Interception of soil water before penetration to deeper roots of M luisana may be the mode of competition between the two species.     

藏东南急尖长苞冷杉群落空间点格局分析及其时空动态

为探究藏东南色季拉山急尖长苞冷杉（Abies georgei var.smithii）群落的空间分布格局及其时空动态,在急尖长苞冷杉纯林中设置1个1 hm²的固定样地,记录样地内所有高度>2 m的急尖长苞冷杉的年龄、胸径、冠幅和坐标等指标。根据径级分布分析种群的径级结构,采用点格局分析方法对种群的空间格局进行分析。结果显示：（1）急尖长苞冷杉的径级分布为倒"J"型,结构稳定,自然更新良好。（2）成树在各个时期和不同尺度上都呈随机分布;幼树在1891年之后的各个时期在小尺度（2 m）上呈聚集分布,1981-2012年在1-41 m尺度上陆续出现聚集分布;幼苗在1861-1920年在小尺度（1-2 m）上呈聚集分布,1951-1980年出现聚集分布的尺度范围最大（1-35 m、41 m和48 m）。（3）在不同时期中,不同年龄组树木之间几乎都呈正关联或无关联;成树和幼树随时间推移从无关联到在各个尺度上陆续出现正关联的变化;成树和幼苗随时间呈无关联、中大尺度（32-35 m、44 m和49 m）正关联和无关联的变化;幼苗和幼树呈现从无关联转变为小尺度（8 m）正关联和小、中尺度（1-30 m）正关联的变化。研究结果表明：成树在小尺度上对幼龄树存在一定的抑制,但在较大尺度是促进的。1951-1980年样地中的幼苗数量大量增加,随后30年幼树的数量因幼苗长大也有增加,这在一定程度上增大了竞争。但总体来看急尖长苞冷杉群落在演替过程中具有种内促进作用。     

Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

Exotic Grass Competition in Suppressing Native Shrubland Re-establishment

Disturbance of coastal sage scrub in southern California has led to extensive displacement of native shrubs by exotic annual grasses. The initial conversion from shrubland to exotic grassland is typically associated with disturbance caused by intense grazing, high fire frequency, or mechanical vegetation removal. While native shrubs have been shown to recolonize annual grasslands under some conditions, other annual grasslands are persistent and show no evidence of shrub recolonization. This study examined the mechanisms by which annual grasses may exclude native shrubs and persist after release from disturbance. Grass density was manipulated in experimental plots to achieve a series of prescribed densities. Artemisia californica, a dominant native shrub, was seeded or planted into the plots and responses to the grass density treatments were measured over two growing seasons. A. californica germination, first season growth, and survival were all negatively related to the density of neighboring annual grasses. The most probable mechanism underlying the reduction of first season growth and survival was depletion of soil water by the grasses. The effects of the grasses on A. californica were no longer significant in the second season. The results of this study indicate that Mediterranean annual grasses reduce recruitment and can persist by inhibiting post-disturbance establishment of A. californica from seed. Although succession alone may not return disturbed annual grasslands to their former shrubland composition, the results suggest that restoration can be achieved by using container plantings or grass removal followed by seeding.     

濒危植物狭果秤锤树群落内主要树种的空间分布格局和关联性

为了解狭果秤锤树(Sinojackia rehderiana)群落的物种共存机制,研究了群落0.5hm~2样地内5主要树种的空间分布格局和种间关联性。结果表明,香樟(Cinnamomum camphora)、紫弹树(Celtis biondii)处于群落上层,尾叶冬青(Ilex wilsonii)在群落中层,下层以狭果秤锤树、瓜木(Alangium platanifolium)为主。狭果秤锤树、香樟、尾叶冬青、瓜木在小尺度上均为聚集分布,随着尺度的增大呈现出均匀分布或随机分布的趋势;紫弹树在全部尺度上为随机分布。尾叶冬青、瓜木在较小尺度上与狭果秤锤树的空间分布呈负相关,而香樟、紫弹树与狭果秤锤树则在较大尺度上呈负相关性,香樟、紫弹树、尾叶冬青、瓜木两两之间均无明显空间相关性。因此,推断密度制约和种子扩散限制对狭果秤锤树群落空间格局形成起作用,生境异质性的影响没有明显表现出来,群落尚未达到稳定状态。     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Effects of fire on the vegetation of a lowland heathland in North-western Italy

This study focuses on the effect of fire on lowland heathlands at the extreme southern edge of their European distribution (Vauda Nature Reserve, NW Italy). Forty-nine plots (50 m radius) were surveyed between 1999 and 2006. Each year, fire occurrences were recorded and per cent cover of four vegetation types (grassland, heath, low shrubland, and tall shrubland) was estimated in each plot. Vascular plant species richness was also recorded in 255, 1 m² quadrats. After a fire, grassland vegetation expanded, but then declined rapidly as heath and shrubland recovered: 7 years after a fire, tall shrubland encroached on to more than 40% of the plots, and grassland declined from 50% to 20% cover. Between 1999 and 2006, Betula pendula shrubland greatly expanded, while grassland decreased over most of the Reserve, even where fire frequency was high. Tall shrubland had low plant diversity and was dominated by widespread species of lower conservation value. By contrast, early successional vegetation (grassland and low shrubland) had higher richness and more narrowly distributed species, indication that the development of tall shrubland causes significant species loss in the heathland. Italian lowland heathlands are characterized by high rates of shrubland encroachment that threatens both habitat and species diversity. Burning frequencies of once in 3–6 years seem appropriate in this habitat, but burning alone might not suffice without actions to increase herbivore grazing.     

Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Effects of an African grass invasion on Hawaiian shrubland nitrogen biogeochemistry

African perennial C₄ grasses are highly successful invaders in Hawaiian ecosystems. We examined the effects of African molasses grass (Melinis minutiflora Beauv.) on Hawaiian shrubland nitrogen (N) dynamics without the influence of fire disturbance. Vegetation tissue carbon and nitrogen chemistry, soil inorganic N pools, net N mineralization rates, and total soil N were studied in three adjacent areas: a monospecificMelinis grassland, a mixed grass/shrubland mosaic, and an un-invaded shrubland.Melinis plots within the mosaic area exhibited the largest inorganic N pools and fastest net N mineralization rates, but were temporally variable with grass phenology. Un-invaded shrubland plots contained the smallest inorganic N pools and lowest net N mineralization rates. Grass foliar C:N and litter C:N were lower than those of common shrubland species, providing one possible link between species and ecosystem N dynamics at this site. The combined effects of N cycle modification, successful light competition, and fire-cycle enhancement make the invasion ofMelinis a significant perturbation to Hawaiian shrubland ecosystem function and successional dynamics. ei]Section editor H Lambers     

Structure and spatial patterns of trees in old-growth northern hardwood and mixed forests of northern Maine

Stand structure including spatial patterns was studied in northern hardwood and mixed forest types in the 2000-ha old-growth Big Reed Forest Reserve in northern Maine using complete stem mapping, dendrochronology, and spatial analyses on 0.5 plots. The inclusion of saplings, dead wood, age distributions, spatial pattern, and interactions provided some idea of underlying processes and temporal change. Structural characteristics were most comparable to spruce-northern hardwood forests of northern New England and New York, and most characteristics matched expected patterns for old-growth forests of the region.Results indicated smaller maximum-tree sizes, lower basal areas (26–34 ) and downed-wood volumes (29–64), higher densities (475–649), but similar species longevities compared to other mesic old-growth forests further south and in the Lake States. The stands were dominated by very shade-tolerant tree species, including Fagus grandifolia Ehrh., Acer saccharum Marsh.,Picea rubens Sarg. and Abies balsamea(L.) Mill, with each species found in many crown positions and age and size classes. The sapling layer was dominated by Fagus grandifolia followed by Picea rubens. Most species had reverse-J shaped diameter distributions, but age distributions were indicative of synchronous, episodic recruitment. In most plots, Acer saccharum diameter distributions were skewed towards the mid-larger size classes. Lack of young and small Acer saccharum stems suggested change in forest composition towards Fagus grandifolia dominance. Most species formed small-scale clusters (15 ) perhaps in response to small gap disturbances. Snags were the dominant dead wood type and were randomly to regularly distributed in most plots. Logfall directions were unrelated to hurricane paths. Recent small-scale disturbance events and topographic position appear to be important in explaining current structure and dynamics of the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. The continued effects of beech bark disease had a greater effect on hardwood plots, whereas a recent spruce budworm outbreak had a greater effect on plots with higher conifer density.The dominance of very shade tolerant tree species in small-scale clusters, and randomly distributed snags rather than clustered uproots were indicative of the prevalence of small scale gap disturbance regimes in the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. Varying topographic position may allow for slight changes in disturbance regime leading to consequent variation in structure and dynamics. H1, a more open plot on upper exposed slopes, had distinctly different characteristics such as lower live and dead tree and sapling densities than the other plots, but more uprooted trees and Acer saccharum saplings. Such small scale gap disturbance regimes operating on an episodic basis, and effects of slight variations in this regime on stand composition and structure have significant implications for silvicultural interventions and management of these forest types.     

Consistent pattern of habitat and species selection by post-dispersal seed predators in a Mediterranean mosaic landscape

There is still little information on effects of habitat degradation on post-dispersal seed predation at the landscape scale. The aim of this study was to determine the influence of habitat degradation and seed species on the variability of post-dispersal seed-predation rate. Experimental seed removal was investigated in six Mediterranean woody plant species, four trees (Pinus sylvestris, Quercus ilex, Acer opalus ssp. granatense, and Sorbus aria) and two shrubs (Berberis vulgaris and Crataegus monogyna), in an extensively used mosaic landscape on the Sierra Nevada massif (SE Spain). Seed depots were distributed over 2 years in five differently degraded landscape units, each one with three plots: shrubland; native forest; and dense, cleared and fenced reforestation stands. Predation was the highest in native forest, shrubland, and fenced reforestation, and the lowest in dense and cleared reforestation stands, being partially due to a positive correlation between shrub cover and post-dispersal seed predation. However, the main factors driving post-dispersal seed predation were intrinsic to seeds, as species preference explained most of the variance in our model for predation. The plant-species ranking was Quercus > Pinus > Sorbus > Berberis > Acer > Crataegus, the dominant tree species being the most depredated. These findings are novel because they suggest for the first time that species-selection patterns by post-dispersal seed predators tended to remain constant through both study years in all habitats comprising a mosaic landscape, whether native forest, reforestation stands or successional shrubland.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Subdominant species distribution in microsites around two life forms at a desert grassland‐shrubland transition zone

Question: In the same landscape context — at a desert grassland‐shrubland transition zone, how does subdominant plant abundance vary in microsites around dominant grasses and shrubs? Location: Sevilleta LTER, New Mexico, USA (34°21’N; 106°53’W; 1650 m a.s.l.). Methods: We compared the distribution of subdominant plants in canopy, canopy edge and interspace microsites around individual shrubs (Larrea tridentata) and grasses (Bouteloua eriopoda) at a transition zone that has been encroached by shrubs within the past 50 ‐ 100 a. Plots of variable size according to microsite type and dominant plant size were sampled. Results: Subdominant abundance was higher in microsites around L. tridentata shrubs than in microsites around B. eriopoda. Furthermore, differences in species abundance and composition were higher among microsites around grasses than among microsites around shrubs. The distribution of subdominants was mostly explained by their phenological characteristics, which indicates the importance of temporal variation in resources to their persistence. Conclusions: This study of coexistence patterns around dominants revealed ecological contrasts between two dominant life forms, but other factors (such as disturbances) have to be taken into consideration to evaluate landscape‐scale diversity.     

Spatial distribution of Eurya japonica in an old-growth evergreen broad-leaved forest,SW Japan

Abstract. The spatial distribution of Eurya japonica, an evergreen understorey species, was studied on a 4-ha permanent plot in an old-growth, temperate, evergreen broad-leaved forest in southwestern Japan. The pattern of spatial dispersion varied with size class, the environmental heterogeneity produced by tree-fall gaps and microtopography and the scale of observation. Seedlings (0.05 to < 0.5 m high) had the most clumped distributions, followed by saplings (0.5 to 1.3 m high) and adults (> 1.3 m). Individuals were more clumped at the smallest grid size, 1.56m × 1.56 m, and became less clumped as grid size increased for all size classes. On grids of less than 6.25 m × 6.25 m, no spatial correlations were found between seedlings and larger size classes; positive spatial correlations were found between saplings and adults at this scale. Abundant seedlings appeared in tree-fall gaps formed by uprooted trees, but seedlings were frequent on ridge tops and upper slopes. Saplings and adults also occurred on upper slopes, but their spatial distribution had little relation to current tree-fall gaps. The spatial distribution of E. japonica individuals was largely attributed to the distribution of tree-fall gaps and the differential responses to microtopographic variation by individuals at different size classes.     

北京山区栓皮栎林优势种群点格局分析

采用点格局分析方法对北京山区栓皮栎林主要种群和不同发育阶段种群的空间分布格局及相互关系进行分析。结果表明:(1)群落物种组成简单,栓皮栎和油松是群落的优势种群,优势种群以中径木为主,缺乏可供更新的幼苗,种群呈现出衰退趋势;(2)在研究尺度内,栓皮栎、油松单种的空间分布格局以聚集型分布为主,随着尺度的增大呈现出随机分布和均匀分布的趋势;栓皮栎和油松呈现显著负相关;(3)优势种群不同发育阶段的空间分布格局与研究尺度紧密相关,栓皮栎小树和中树以聚集型分布为主、大树呈随机分布,而油松不同发育阶段均以随机分布为主;(4)栓皮栎、油松种群不同发育阶段之间的空间关联主要发生在小尺度范围内,小树和中树以正关联为主,而中树和大树、小树和大树以负关联为主;随着研究尺度的增大趋于无关联。     

Distribution,species diversity and life-form spectra of plant communities along an altitudinal gradient in the northern slopes of Qilianshan Mountains,Gansu, China

We studied the distribution pattern, species diversity and life-formspectra of plant communities along an altitudinal gradient in the mid-sectionofthe northern slopes of Qilianshan Mountains by means of multivariate analyses.Two data sets (167 species × 75 plots, 10 environmental variables ×75 plots), originated from the fieldworks in 1998–1999, were subjected toTWINSPAN and DCCA, resulting in 8 major plant communities: 1)Asterothamnus centraliasiaticus–Halogetonarachnoideus desert grassland on azonal substrates from 1450 to 1600m and 2) zonal Reaumuria soogorica desertgrassland on gravels from 1470 to 1900 m; 3) Stipaprzewalskii–Stipa purpurea montane grassland from 2200 to 2900m; 4) Polygonum viviparum alpine grasslandfrom 2900 to 3700 m; 5) Caraganastenophylla–Ajaniafruticulosa dry-warm shrubland from 2350 to 2800 m; 6)Sabina przewalskii mid-wet warm forest from 2700 to 3300m; 7) Picea crassifolia cold coniferousforestfrom 2450 to 3200 m; 8) Caragana jubatawet-cold alpine shrubland from 3100 to 3700 m. Species diversityand species richness of both grasslands and forests peaked at the intermediateportion of the elevational gradient. Evenness might be strongly influenced byeither biotic or abiotic factors at a local scale, while seems quiteindependentof an elevational gradient at landscape scales. Beta-diversity decreased from1500 to 3700 m, indicating that species turnover declined withincreased elevation. Both richness of life-form and total species richness in agiven altitudinal belt (gamma-diversity) peaked at intermediate elevations,while relative species richness of different life-form varied differently alongthe altitudinal gradient.