Stability of the spatio-temporal distribution and niche overlap in neotropical earthworm assemblages

The spatial distribution of soil invertebrates is aggregated with high-density patches alternating with low-density zones. A high degree of spatio-temporal organization generally exists with identified patches of specific species assemblages, in which species coexist according to assembly rules related to competitive mechanisms for spatial and trophic resources occur. However, these issues have seldom been addressed. The spatio-temporal structure of a native earthworm community in a natural savanna and a grass–legume pasture in the Colombian “Llanos” was studied during a 2-year-period. A spatially explicit sampling design (regular grid) was used to discern the distribution pattern of species assemblages in both systems. Earthworms were collected from small soil pits at three different sampling dates. Data collected from 1 m² soil monoliths were also used in the present study. Data were analyzed with the partial triadic analysis (PTA) and correlograms, while niche overlap was computed with the Pianka index. The PTA and correlogram analysis revealed that earthworm communities displayed a similar stable spatial structure in both systems during the 2-year study period. An alternation of population patches where different species' assemblages dominated was common to all sampling dates. The medium-sized Andiodrilus sp. and Glossodrilus sp. exhibited a clear spatial opposition in natural savanna and the grass–legume pasture for the duration of the study. The Pianka index showed a high degree of niche overlapping in several dimensions (vertical distribution, seasonality of population density) between both species. The inclusion of space-time data analysis tools as the PTA and the use of classical ecological indices (Pianka) in soil ecology studies may improve our knowledge of earthworm assemblages' dynamics.     

Stability of the spatio-temporal distribution and niche overlap in neotropical earthworm assemblages

The spatial distribution of soil invertebrates is aggregated with high-density patches alternating with low-density zones. A high degree of spatio-temporal organization generally exists with identified patches of specific species assemblages, in which species coexist according to assembly rules related to competitive mechanisms for spatial and trophic resources occur. However, these issues have seldom been addressed. The spatio-temporal structure of a native earthworm community in a natural savanna and a grass–legume pasture in the Colombian “Llanos” was studied during a 2-year-period. A spatially explicit sampling design (regular grid) was used to discern the distribution pattern of species assemblages in both systems. Earthworms were collected from small soil pits at three different sampling dates. Data collected from 1 m² soil monoliths were also used in the present study. Data were analyzed with the partial triadic analysis (PTA) and correlograms, while niche overlap was computed with the Pianka index. The PTA and correlogram analysis revealed that earthworm communities displayed a similar stable spatial structure in both systems during the 2-year study period. An alternation of population patches where different species' assemblages dominated was common to all sampling dates. The medium-sized Andiodrilus sp. and Glossodrilus sp. exhibited a clear spatial opposition in natural savanna and the grass–legume pasture for the duration of the study. The Pianka index showed a high degree of niche overlapping in several dimensions (vertical distribution, seasonality of population density) between both species. The inclusion of space-time data analysis tools as the PTA and the use of classical ecological indices (Pianka) in soil ecology studies may improve our knowledge of earthworm assemblages' dynamics.     

Soil environmental heterogeneity allows spatial co‐occurrence of competitor earthworm species in a gallery forest of the Colombian ‘Llanos’

Disentangling how communities of soil organisms are deterministically structured by abiotic and biotic factors is of utmost relevance, and few data sets on co‐occurrence patterns exist in soil ecology compared to other disciplines. In this study, we assessed species spatial co‐occurrence and niche overlap together with the heterogeneity of selected soil properties in a gallery forest (GF) of the Colombian Llanos. We used null‐model analysis to test for non‐random patterns of species co‐occurrence and body size in assemblages of earthworms and whether the pattern observed was the result of environmental heterogeneity or biotic processes structuring the community at small scales by means of co‐inertia analysis (CoIA). The results showed that earthworm species co‐occurred more frequently than expected by chance at short distances, and CoIA highlighted a significant specific relationship between earthworm species and soil variables. The effect of soil environmental heterogeneity on one litter‐feeding species but also the impact of soil‐feeding species on soil physical properties was revealed. Correlogram analysis on the first axis extracted in the CoIA showed the scale of the common structure shared by the fauna and soil variable tables. The earthworm community was not deterministically structured by competition and co‐occurrence of competing species was facilitated by soil environmental heterogeneity at small scales in the GF. Our results agreed with the coexistence aggregation model which suggests that spatial aggregation of competitors at patchily distributed resources (environment) can facilitate species coexistence.     

The importance of landscape and habitat properties in explaining instantaneous and long‐term distributions of large branchiopods in subtropical temporary pans

1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long‐term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long‐term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.     

Soil heterogeneity and community composition jointly influence grassland biomass

Question: Does the spatial pattern of nutrient supply modify community biomass responses to changes in both species composition and richness? Location: Duke University Phytotron (Durham, North Carolina, USA). Methods: We conducted a microcosm experiment to evaluate individual plant and whole community responses to species richness, species composition and soil nutrient heterogeneity. The experiment consisted of seven levels of species composition (all possible combinations of Lolium perenne, Poa pratensis and Plantago lanceolata) crossed with three levels of soil nutrient distribution (homogeneous, heterogeneous‐up, and heterogeneous‐down, where up and down indicates the location of a nutrient patch in either the upper or the lower half of the soil column, respectively). Results: Communities containing Plantago and Lolium responded to nutrient heterogeneity by increasing above‐ and below‐ground biomass. Nutrient heterogeneity also increased size inequalities among individuals of these species. Significant species composition X nutrient heterogeneity interactions on community biomass and individual size inequality were observed when nutrient patches were located in the upper 10 cm of the soil columns. However, root proliferation in nutrient patches was equivalent regardless of the vertical placement of the patch. Conclusions: Our results suggest that nutrient heterogeneity may interact with plant species composition to determine community biomass, and that small‐scale vertical differences in the location of nutrient patches affect individual and community responses to this heterogeneity.     

Spatial and temporal community structure of desmids on a small spatial scale

Besides spatial heterogeneity, another important component of the diversity of protist communities is the variation in species assemblages through time. Despite its importance, temporal turnover of benthic communities has been studied to a lesser extent than spatial heterogeneity has. In this study, we examine the desmid assemblages on small spatial scale in relation to the spatial, temporal, and environmental parameters. The samples were collected within two different types of peatland localities in the Czech Republic over 3 years. The differences in species composition between samples were mainly correlated with the geographic distance, while the effects of the environmental and temporal variables were much weaker. Since the spatial heterogeneity of the assemblages was not induced by the variation of the environmental factors or by the restricted dispersal ability at such a small spatial scale, we assume that both the temporal stability and strong spatial autocorrelation might have been the result of a priority effect, with subsequent monopolization of resources. Stochasticity in colonization can introduce noise into the match between community composition and environmental conditions, which may result in stronger effect of the spatial parameters on the community structure.     

Effects of Soil Nutrient Heterogeneity and Earthworms on Aboveground Biomass of Experimental Plant Communities

Soil nutrients are commonly heterogeneously distributed and earthworms are one of the most common soil organisms. While effects of both soil nutrient heterogeneity and earthworms have been well studied, their interactive effect on plant community productivity has rarely been tested. In a greenhouse experiment, we constructed experimental plant communities by sowing seed mixtures of four grasses, two legumes and two forbs in either a heterogeneous soil consisting of low and high nutrient soil patches or a homogeneous soil where the low and high nutrient soil patches were evenly mixed. The earthworm Eisenia fetida was either added to these soils or not. Aboveground biomass of the whole communities, grasses and legumes did not differ between the homogeneous and heterogeneous soils or between the soils with and without earthworms. However, soil nutrient heterogeneity reduced aboveground biomass of forbs, and such an effect did not interact with earthworms. In response to soil heterogeneity and earthworms, biomass ratio of the three functional groups showed similar patterns as that of their biomass. At the patch level, aboveground biomass of the whole community, grasses and legumes were greater in the high than in the low nutrient soil patches within the heterogeneous soil. A similar pattern was found for the forbs, but this was only true in the absence of earthworms. Our results suggest that soil nutrient heterogeneity and earthworms may not influence aboveground biomass of plant communities, despite the fact that they may modify the growth of certain plant functional groups within the community.     

Spatial Patchiness of Litter, Nutrients and Macroinvertebrates during Secondary Succession in a Tropical Montane Cloud Forest in Mexico

We test the hypothesis that secondary succession in Tropical Montane Cloud Forest (TMCF) in Mexico is accompanied by an increase in the spatial structuring of litter resources, soil nutrient concentrations and the soil macroinvertebrate community at a within-plot scale (5–25 m). This increased spatial structuring is expected because secondary succession in these forests is associated with an increase in the diversity of trees that dominate the canopy. If each tree species generates a particular soil environment under its canopy, then under a diverse tree community, soil properties will be spatially very heterogeneous. Tree censuses and grid sampling were performed in four successional stages of a secondary chronosequence of TMCF. Variography was used to analyse spatial patterns in continuous variables such as nutrient concentrations, while Spatial Analysis by Distance Indices (SADIE) was applied to determine patchiness in the distribution of soil macroinvertebrate taxa. Secondary succession was found to be accompanied by the predicted increase in the spatial structuring of litter resources and the macroinvertebrate community at the within-plot scale. Spatial patterns in the macroinvertebrate community only became evident for all taxa in the oldest forest (100 years old). Patches with low Ca and Mg concentrations in early successional soils were associated with patches where pine litter was most abundant while those with low P concentrations in late successional stages were associated with patches where oak litter was most abundant. Results suggest that anthropogenic disturbance aboveground promotes a more homogeneous resource environment in the surface soil, which compared to older forests, sustains a less diverse and less spatially structured macroinvertebrate community.     

Spatial gradients in species diversity of microscopic animals: the case of bdelloid rotifers at high altitude

Aim Organisms smaller than 2 mm appear not to follow the spatial patterns in richness and diversity commonly observed in macroscopic organisms. We describe spatial patterns in species diversity in a group of microscopic organisms, bdelloid rotifers, living in moss and lichen patches, in order to test the hypotheses of no relationship between species richness and composition and spatial gradients, suggested by previously published patterns in microscopic organisms. Location Moss and lichen patches as habitats for bdelloids, on high‐elevation peaks at altitudes between 2984 and 4527 m a.s.l. across the Italian, French and Swiss Alps, with distances among sample sites ranging from 1 m to 420 km, in comparison with lower‐elevation samples at altitudes from 850 to 1810 m a.s.l. Methods We sampled species assemblages of bdelloid rotifers living in isolated moss and lichen patches in 47 sites. We described the observed α, β and γ diversities; the heterogeneity of species assemblages; and the estimated number of species (incidence‐based coverage estimator). Patterns in species distribution were analysed at three different levels: (1) habitat, comparing species richness on moss and lichen substrates, testing differences in α diversity and heterogeneity (anova ), species composition (analysis of similarities test), and γ diversity (rarefaction curves); (2) altitude, comparing the observed richness with previously published data from locations well below 2000 m; and (3) distances between sites, correlating the matrix of Jaccard dissimilarities and the matrix of geographical distances with a Mantel test. Results Both species richness and species composition of bdelloid rotifers differed significantly between mosses and lichens at high elevations, but no difference was found in the heterogeneity of species assemblages. Alpha diversity was significantly lower at high‐elevation than at low‐elevation sites, but the estimated number of species was not reduced when compared with sites at low elevations. Geographical distance between sites had no effect on species composition in either mosses or lichens. The distribution of species was highly heterogeneous, with a low similarity among assemblages. Main conclusions As expected, bdelloids appear to occupy habitats selectively. The altitudinal gradient in species richness for bdelloid rotifers is limited to a decrease in α diversity only; such a decrease is not caused by a lower number of species (low γ diversity) being able to tolerate harsh conditions, and high‐altitude species are not a subset of species living at lower elevations. The observed values of α, β and γ diversity at high altitudes in the Alps are compatible with the scenario of a very low number of available propagules because of the low density of patches of favourable habitat. Our results suggest that the geographical distribution of animals, and therefore biodiversity patterns, may be strongly influenced by animal size, as small organisms such as bdelloids appear to show spatial patterns that differ from those known in larger animals. Differences in body size should be taken into account carefully in future studies of biodiversity patterns.     

Plant–soil relationships in fragments of Mediterranean snow-beds: ecological and conservation implications

We investigated the spatial structures of soil properties and snow-bed vegetation, and their relationships, in southern Italy. We analyzed data on 26 plant species and 10 soil traits from adjacent 1 × 1 m plots in two snow-bed patches. Measures of spatial autocorrelation revealed striking spatial structures for plant cover and soil properties at both sites. Bivariate statistics and Mantel tests highlighted a significant correlation between spatial patterns of plants and soil in the study sites. Canonical correspondence analysis related such relationships to an ecological gradient connecting soil properties and plant assemblages in this unusual ecological context. Among the variables significantly related to plant patterns is the soil organic matter, which is recognized as being sensitive to global warming. Our analyses suggest that soil dynamics due to increasing temperature may promote the replacement of species typical of southern snow-bed ecosystems by more mesophilous plants.     

Small‐Scale Distribution of Aquatic Macroinvertebrates in Two Spring Fens with Different Groundwater Chemistry

We examined responses of macroinvertebrate assemblages to environmental and temporal variations along spring source‐spring brook transects in two fen habitats, sharply differing in groundwater chemistry, and compared the patterns among individual taxonomical groups. We hypothesised a different importance of environmental heterogeneity and seasonal changes primarily linked to strong tufa precipitation, which causes stronger environmental filtering in the calcareous fen. In concordance, we observed that assemblages of the more homogenous calcareous fen primarily changed over time, due to seasonal shifts in source availability and favourable conditions. Their spatial distribution was determined by the amount of CPOM, tufa crusts and temperature variation, but a substantial part of the assemblage exhibited spatial uniformity (Plecoptera, Clitellata, and especially Trichoptera and Diptera). The assemblages of the more heterogeneous Sphagnum ‐fen were primarily driven by water pH and substrate and the season was a notably weaker predictor. We found that different macroinvertebrate groups can display various responses to the measured variables shaping the overall pattern obtained based on the whole community. Further, greater environmental heterogeneity can result in temporally stable species distribution patterns even at very small spatial scales within a single site. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The quest for a null model for macroecological patterns: geometry of species distributions at multiple spatial scales

There have been several attempts to build a unified framework for macroecological patterns. However, these have mostly been based either on questionable assumptions or have had to be parameterized to obtain realistic predictions. Here, we propose a new model explicitly considering patterns of aggregated species distributions on multiple spatial scales, the property which lies behind all spatial macroecological patterns, using the idea we term 'generalized fractals'. Species' spatial distributions were modelled by a random hierarchical process in which the original 'habitat' patches were randomly replaced by sets of smaller patches nested within them, and the statistical properties of modelled species assemblages were compared with macroecological patterns in observed bird data. Without parameterization based on observed patterns, this simple model predicts realistic patterns of species abundance, distribution and diversity, including fractal-like spatial distributions, the frequency distribution of species occupancies/abundances and the species–area relationship. Although observed macroecological patterns may differ in some quantitative properties, our concept of random hierarchical aggregation can be considered as an appropriate null model of fundamental macroecological patterns which can potentially be modified to accommodate ecologically important variables.     

The influence of soil depth on plant species response to grazing within a semi-arid savanna

Grassland patches within a semi-arid savanna were evaluated over 45-years for (1) local temporal dynamics of basal area for five dominant grass species within long-term heavily grazed and ungrazed treatments, (2) the influence of soil depth (resource availability) on vegetation dynamics, and (3) the applicability of community-level grazing response groups over fine-scale patterns of soil heterogeneity. Temporal patterns in species composition and basal area were dependent upon soil depth. In the heavy grazed treatment, Hilaria belangeri dominated deep soils while Erioneuron pilosum and Bouteloua trifida were restricted to shallow soils. In the ungrazed treatment, removal of grazing resulted in successional changes that were significantly different across soil depths. After 45 years without grazing, Eriochloa sericea was most abundant on deep soils while Bouteloua curtipendula was more abundant on intermediate and shallow soils. Community-level functional groups that are based on grazing were not appropriate when multiple pattern-driving variables were considered across multiple scales indicating that functional groups should only be applied to certain processes at specific scales. Within the ungrazed treatments, variable soil depths have resulted in a shifting mosaic in time and space where early- and late-successional species co-exist continuously but spatially separated within the community. In the heavily grazed treatment, species are somewhat spatially arranged by soil depths, but much of the inherent heterogeneity is eliminated and species composition is dominated by the three grazing-resistant short-grasses. Broad scale successional changes may appear linear and predictable while at finer scales, the same changes may be described as non-linear and dependent upon soil depth resulting in thresholds that are partially explained by weather patterns, seed bank limitations and competitive inhibitions.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Disturbance alters habitat isolation's effect on biodiversity in aquatic microcosms

Isolated habitats generally have fewer species at local spatial scales than more connected habitats. However, over larger spatial scales, the response of species richness to variation in the degree of isolation is variable. Here, we hypothesized that the effects of habitat isolation on patterns of regional level species richness may depend at least in part on the level of disturbances those habitats receive. We tested this hypothesis in a microcosm experiment using an aquatic community consisting of container dwelling protists and rotifers by manipulating disturbance and dispersal to experimental regions factorially. In disturbed regions, regional species richness was lower in regions with isolated patches compared to regions where patches were experimentally connected by dispersal. A likely mechanism for this result is that dispersal from adjacent undisturbed local patches allowed disturbance-intolerant species a temporary refugia, thereby allowing regional coexistence of disturbance-tolerant and intolerant species. In contrast, without disturbances (and thus no temporal heterogeneity) it is likely that dispersal homogenized communities, leading to overall lower richness with higher dispersal. Our results emphasize the importance of simultaneously considering multiple limiting factors, disturbance and dispersal in this case, as well as the spatial scale of the response, in order to fully understand factors that control biodiversity.     

Small-scale spatial variability in intertidal and subtidal turfing algal assemblages and the temporal generality of these patterns

Spatial and temporal variation in patterns of distribution and abundance of algal assemblages is large and often occurs at extremely small spatial and temporal scales. Despite this, few studies investigate interactions between these scales, that is, how patterns of spatial variation change through time. This study investigated a number of scales of spatial variation (from tens of centimetres to kilometres) in assemblages of intertidal and subtidal turfing algae. Significant differences were found in the composition and abundances of species in assemblages of turf at all spatial scales tested. Much of the variation among assemblages could, however, be explained at the scale of quadrats (tens of centimetres apart) (27±1.4 (SE)% of dissimilarity) with an additional 7±1.2% explained at the scale of sites (tens of metres apart) and 10±1.5% at the scale of locations (kilometres apart). Although the greatest dissimilarity in assemblages occurred at the scale of habitats, this accounted for a relatively small proportion of the overall variation in assemblages. These patterns were consistent through time, that is, at each sampling time the spatial scale explaining the greatest proportion of variation in assemblages was replicate quadrats separated by tens of centimetres. These patterns appear to be due to small-scale variation in patterns of distribution and abundances of the individual species that comprise turfing algal assemblages. The results of this experiment suggest that large scale processes have less effect on patterns of variability of algal assemblages than those occurring on relatively smaller spatial scales and that small-scale spatial variation should not be considered as simply “noise”.     

古尔班通古特沙漠土壤水分与化学性质的空间分布

区域尺度上,气象和水文状况是影响植被分布格局和土壤性状的主要因子,当然,局部的地貌特征及植被分布对土壤性质的影响也是不能忽视的。以古尔班通古特沙漠为研究对象,对其南缘至腹地约25 km的土壤水分与化学性质的空间分布及其相互关系进行研究,结果显示:水平方向上,从沙漠南缘至腹地,土壤水分在丘底呈减小趋势,尤其是距边缘10 km以外,且在距边缘6—7 km出现1个峰值。同时土壤pH值和电导率的大小以及有机碳,全氮,有效氮,全磷和有效磷含量与土壤水分的变化状况相似,在丘底呈减小趋势,而在丘坡和丘顶上均呈波动分布。这说明丘底土壤性质的空间格局受到气象和水文状况的显著影响,而丘坡和丘顶土壤性质并没有受到气象和水文状况的显著影响,这种波动分布主要由采样点的设置及植物的分布状况所引起。丘底土壤水分及化学性质均显著高于丘坡和丘顶,这说明地貌特征对土壤性质的空间分布有显著影响。垂直方向上,不同深度土层土壤性质也存在显著差异,土壤水分,pH值和电导率随着土层深度增加而增加,而土壤养分随土层深度增加而减小。相关分析结果表明:仅4、5月份的土壤含水量及年平均土壤含水量与丘底土壤化学性质显著正相关,这可能由于冬季积雪的融化导致了土壤水分在4、5月份的聚集比较明显,使土壤水分对土壤化学性质发挥作用。总之,土壤空间异质性是由气象和水文以及地貌特征共同影响的,而土壤空间异质性的变异进而影响到植被的分布,植被的分布反过来又影响土壤的空间分布,因此土壤的空间分布和植被的空间分布是相互影响,共同作用的。