Assessing topographic patterns in moisture use and stress using a water balance approach

Through its control on soil moisture patterns, topography’s role in influencing forest composition is widely recognized. This study addresses shortcomings in traditional moisture indices by employing a water balance approach, incorporating topographic and edaphic variability to assess fine-scale moisture demand and moisture availability. Using GIS and readily available data, evapotranspiration and moisture stress are modeled at a fine spatial scale at two study areas in the US (Ohio and North Carolina). Model results are compared to field-based soil moisture measurements throughout the growing season. A strong topographic pattern of moisture utilization and demand is uncovered, with highest rates of evapotranspiration found on south-facing slopes, followed by ridges, valleys, and north-facing slopes. South-facing slopes and ridges also experience highest moisture deficit. Overall higher rates of evapotranspiration are observed at the Ohio site, though deficit is slightly lower. Based on a comparison between modeled and measured soil moisture, utilization and recharge trends were captured well in terms of both magnitude and timing. Topographically controlled drainage patterns appear to have little influence on soil moisture patterns during the growing season. In addition to its ability to accurately capture patterns of soil moisture in both high-relief and moderate-relief environments, a water balance approach offers numerous advantages over traditional moisture indices. It assesses moisture availability and utilization in absolute terms, using readily available data and widely used GIS software. Results are directly comparable across sites, and although output is created at a fine-scale, the method is applicable for larger geographic areas. Since it incorporates topography, available water capacity, and climatic variables, the model is able to directly assess the potential response of vegetation to climate change.     

Ecosystem-scale spatial heterogeneity of stable isotopes of soil nitrogen in African savannas

Soil ¹⁵N is a natural tracer of nitrogen (N) cycling. Its spatial distribution is a good indicator of processes that are critical to N cycling and of their controlling factors integrated both in time and space. The spatial distribution of soil δ¹⁵N and its underlying drivers at sub-kilometer scales are rarely investigated. This study utilizes two sites (dry vs. wet) from a megatransect in southern Africa encompassing locations with similar soil substrate but different rainfall and vegetation, to explore the effects of soil moisture and vegetation distribution on ecosystem-scale patterns of soil δ¹⁵N. A 300-m long transect was set up at each site and surface soil samples were randomly collected for analyses of δ¹⁵N, %N and nitrate content. At each soil sampling location the presence of grasses, woody plants, Acacia species (potential N fixer) as well as soil moisture levels were recorded. A spatial pattern of soil δ¹⁵N existed at the dry site, but not at the wet site. Woody cover distribution determined the soil δ¹⁵N spatial pattern at ecosystem-scale; however, the two Acacia species did not contribute to the spatial pattern of soil δ¹⁵N. Grass cover was negatively correlated with soil δ¹⁵N at both sites owing to the lower foliar δ¹⁵N values of grasses. Soil moisture did not play a role in the spatial pattern of soil δ¹⁵N at either site. These results suggest that vegetation distribution, directly, and water availability, indirectly, affect the spatial patterns of soil δ¹⁵N through their effects on woody plant and grass distributions.     

Spatial dependence and the relationship of soil organic carbon and soil moisture in the Luquillo Experimental Forest,Puerto Rico

We used geo-spatial statistical techniques to examine the spatial variation and relationship of soil organic carbon (SOC) and soil moisture (SM) in the Luquillo Experimental Forest (LEF), Puerto Rico, in order to test the hypothesis that mountainous terrain introduces spatial autocorrelation and crosscorrelation in ecosystem and soil properties. Soil samples (n = 100) were collected from the LEF in the summer of 1998 and analyzed for SOC, SM, and bulk density (BD). A global positioning system was used to georeference the location of each sampling site. At each site, elevation, slope and aspect were recorded. We calculated the isotropic and anisotropic semivariograms of soil and topographic properties, as well as the cross-variograms between SOC and SM, and between SOC and elevation. Then we used four models (random, linear, spherical and wave/hole) to test the semi-variances of SOC, SM, BD, elevation, slope and aspect for spatial dependence. Our results indicate that all the studied properties except slope angle exhibit spatial dependence within the scale of sampling (200 – 1000 m sampling interval). The spatially structured variance (the variance due to the location of sampling sites) accounted for a large proportion of the sample variance for elevation (99%), BD (90%), SOC (68%), aspect (56%) and SM (44%). The ranges of spatial dependence (the distances within which parameters are spatially dependent) for aspect, SOC, elevation, SM, and BD were 9810 m, 3070 m, 1120 m, 930 m and 430 m, respectively. Cross correlograms indicate that SOC varies closely with elevation and SM depending on the distances between samples. The correlation can shift from positive to negative as the separation distance increases. Larger ranges of spatial dependence of SOC, aspect and elevation indicate that the distribution of SOC in the LEF is determined by a combination of biotic (e.g., litterfall) and abiotic factors (e.g., microclimate and topographic features) related to elevation and aspect. This demonstrates the importance of both elevation and topographic gradients in controlling climate, vegetation distribution and soil properties as well as the associated biogeochemical processes in the LEF.This revised version was published online in May 2005 with corrections to the Cover Date.     

Road verges as invasion corridors? A spatial hierarchical test in an arid ecosystem

Disturbed habitats are often swiftly colonized by alien plant species. Human inhabited areas may act as sources from which such aliens disperse, while road verges have been suggested as corridors facilitating their dispersal. We therefore hypothesized that (i) houses and urban areas are propagule sources from which aliens disperse, and that (ii) road verges act as corridors for their dispersal. We sampled presence and cover of aliens in 20 plots (6 × 25 m) per road at 5-km intervals for four roads, nested within three localities around cities (n = 240). Plots consisted of three adjacent nested transects. Houses (n = 3,349) were mapped within a 5-km radius from plots using topographical maps. Environmental processes as predictors of alien composition differed across spatial levels. At the broadest scale road-surface type, soil type, and competition from indigenous plants were the strongest predictors of alien composition. Within localities disturbance-related variables such as distance from dwellings and urban areas were associated with alien composition, but their effect differed between localities. Within roads, density and proximity of houses was related to higher alien species richness. Plot distance from urban areas, however, was not a significant predictor of alien richness or cover at any of the spatial levels, refuting the corridor hypothesis. Verges hosted but did not facilitate the spread of alien species. The scale dependence and multiplicity of mechanisms explaining alien plant communities found here highlight the importance of considering regional climatic gradients, landscape context and road-verge properties themselves when managing verges. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Topography-mediated controls on local vegetation phenology estimated from MODIS vegetation index

Forest canopy phenology is an important constraint on annual water and carbon budgets, and responds to regional interannual climate variation. In steep terrain, there are complex spatial variations in phenology due to topographic influences on microclimate, community composition, and available soil moisture. In this study, we investigate spatial patterns of phenology in humid temperate forest as a function of topography. Moderate-resolution imaging spectro-radiometer (MODIS) vegetation indices are used to derive local patterns of topography-mediated vegetation phenology using a simple post-processing analysis and a non-linear model fitting. Elevation has the most explanatory power for all phenological variables with a strong linear relationship with mid-day of greenup period, following temperatures lapse rates. However, all other phenological variables show quadratic associations with elevation, reflecting an interaction between topoclimatic patterns of temperature and water availability. Radiation proxies also have significant explanatory power for all phenological variables. Though hillslope position cannot be adequately resolved at the MODIS spatial resolution (250 m) to discern impacts of local drainage conditions, extended periods of greenup/senescence are found to occur in wet years. These findings are strongly supported by previous field measurements at different topographic positions within the study area. The capability of detecting topography-mediated local phenology offers the potential to detect vegetation responses to climate change in mountainous terrain. In addition, the large, local variability of meteorological and edaphic conditions in steep terrain provides a unique opportunity to develop an understanding of canopy response to the interaction of climate and landscape conditions.     

Spatial analysis of selected soil attributes across an alpine topographic/snow gradient

The impact of the topographic/snow gradient on soil processes in alpinetundra on Niwot Ridge of the Colorado Front Range (Rocky mts, USA) was assessedusinggeostatistical modeling and a fractal approach. The mean snow depth, whichmeasured between 1984 and 2000, exhibited a smooth spatial continuity acrossthestudy grid area (550 × 400 meter). Soil color variables showed a nestedstructure that was attributed to a confounded effect of various soil-formingfactors on catenary processes. The spatial structure of texture classesexhibited no spatial structure and was explained by data sparsity,cryoturbation, and biological processes that mask the expected long-distancevariations (i.e., 550-m) of the catenary processes. Organic C, pH, bulkdensity,and soil moisture content showed various degrees of spatial continuity, but allindicated that the topographic/snow gradient is not the only dominantsoil-forming factor in this alpine ecosystem. The estimated fractal dimensionDfor the grid landform and the mean snow depth varied between 1.2 and 1.4,indicating that they vary smoothly with long-range variation. The estimatedDofthe soil variables ranged between 1.6 and 1.8, showing a noisy appearance withshort-range variations. These results strongly suggest that most small andmicro-scale variations in the alpine soil environs resulted from the combinedeffect of cryoturbation, biological activity, parent-material and eoliandeposition, whereas the large-scale variations originated as a result of thetopographic/snow gradient.This revised version was published online in May 2005 with corrections to the Cover Date.     

Multiscale control of vegetation patterns: the case of Doñana (SW Spain)

The early studies about the plant ecology of Doñana carried out at a small scale showed that the main process controlling vegetation composition of the stabilized dunes was soil water availability. However, the extrapolation of this model to larger spatial scales failed to explain observed vegetation patterns. In this work, the vegetation patterns and the processes causing them are studied at a larger scale. Data of topography, soil pH, electrical conductivity, and available iron allowed to distinguish three large geomorphologic zones on the stabilized dunes of the Doñana Biological Reserve which correspond to different dune building episodes. Different dune episodes showed differences in both water table depth and dynamics, which are due to groundwater flow systems of different scale. It is further manifested by differences in shrub composition. The results show that geomorphology controls the vegetation pattern at different scales mediated through water availability. Differences in water availability are due to the connection to groundwater flow systems of contrasted scale. On a small scale (10−10² m), along dune slopes, there is a gradient from dune ridges to slacks, from xerophyte to hygrophyte vegetation types. On a mesoscale (10²−10³ m), there are several dune episodes with variable topographic altitude, dominated by different types of xerophytes. On a regional scale (>10³ m), the discharges of the regional aquifer produce strong environmental and biotic stresses resulting in a mixed community.     

Small-scale effects of historical land use and topography on post-cultural tree species composition in an Alpine valley in southern Switzerland

Investigations of spatial patterns in forest tree species composition are essential in the understanding of landscape dynamics, especially in areas of land-use change. The specific environmental factors controlling the present patterns, however, vary with the scale of observation. In this study we estimated abundance of adult trees and tree regeneration in a Southern Alpine valley in Ticino, Switzerland. We hypothesized that, at the present scale, spatial pattern of post-cultural tree species does not primarily depend on topographic features but responds instead to small-scale variation in historical land use. We used multivariate regression trees to relate species abundances to environmental variables. Species matrices were comprised of single tree species abundance as well as species groups. Groups were formed according to common ecological species requirements with respect to shade tolerance, soil moisture and soil nutrients. Though species variance could only be partially explained, a clear ranking in the relative importance of environmental variables emerged. Tree basal area of formerly cultivated Castanea sativa (Mill.) was the most important factor accounting for up to 50% of species’ variation. Influence of topographic attributes was minor, restricted to profile curvature, and partly contradictory in response. Our results suggest the importance of biotic factors and soil properties for small-scale variation in tree species composition and need for further investigations in the study area on the ecological requirements of tree species in the early growing stage.     

Associations between soil carbon and ecological landscape variables at escalating spatial scales in Florida,USA

The spatial distribution of soil carbon (C) is controlled by ecological processes that evolve and interact over a range of spatial scales across the landscape. The relationships between hydrologic and biotic processes and soil C patterns and spatial behavior are still poorly understood. Our objectives were to (i) identify the appropriate spatial scale to observe soil total C (TC) in a subtropical landscape with pronounced hydrologic and biotic variation, and (ii) investigate the spatial behavior and relationships between TC and ecological landscape variables which aggregate various hydrologic and biotic processes. The study was conducted in Florida, USA, characterized by extreme hydrologic (poorly to excessively drained soils), and vegetation/land use gradients ranging from natural uplands and wetlands to intensively managed forest, agricultural, and urban systems. We used semivariogram and landscape indices to compare the spatial dependence structures of TC and 19 ecological landscape variables, identifying similarities and establishing pattern–process relationships. Soil, hydrologic, and biotic ecological variables mirrored the spatial behavior of TC at fine (few kilometers), and coarse (hundreds of kilometers) spatial scales. Specifically, soil available water capacity resembled the spatial dependence structure of TC at escalating scales, supporting a multi-scale soil hydrology-soil C process–pattern relationship in Florida. Our findings suggest two appropriate scales to observe TC, one at a short range (autocorrelation range of 5.6 km), representing local soil-landscape variation, and another at a longer range (119 km), accounting for regional variation. Moreover, our results provide further guidance to measure ecological variables influencing C dynamics.     

Scaling patterns of biomass and soil properties: an empirical analysis

We argue that studies at multiple scales must necessarilychange the extent of measurements, not just the spacing, in order toeffectivelycapture information regarding processes at multiple scales. We have implementeda multi-scale sampling scheme using transects of 10 cm, 1m, 10 m, 100 m, and 1 km ateach of four sites along an elevational gradient from dry foothills forest toalpine tundra in the Front Range of Colorado; these four sites form anadditional transect of 22 km. Along each of these transects wetookten equally spaced soil cores and measured variables important in determiningboth microbial and plant community structure: soil water content, organicmattercontent, pH, and total soil biomass. With this sampling scheme we are able totreat scale as an independent variable in our analyses, and our data show thatboth particular sites and particular variables can determine whether estimatesof mean values are scale-dependent or not. A geostatistical analysis using allof our data shows common relationships between scales across ecologicallydiverse sites; biomass shows the most complex pattern of distribution acrossscales, as measured by fractal dimension. Our analyses also reveal theinadequacy of several standard geostatistical models when applied to data frommultiple scales of measurement – we recommend the use of the boundedpowerlaw model in such cases. We hypothesize that because biological communitiesmustrespond simultaneously to multiple variables with differing patterns of spatialvariation, the spatial variation of biological communities will be at least ascomplex as the most complex environmental variable at any given site.This revised version was published online in May 2005 with corrections to the Cover Date.     

Estimation and prediction of plant species richness in a mosaic landscape

Traditional agricultural mosaic landscapes are likely to undergo dramatic changes through either intensification or abandonment of land use. Both developmental trends may negatively affect the vascular plant species richness of such landscapes. Therefore, sustainable land-use systems need to be developed to maintain and re-establish species richness at various spatial scales. To evaluate the sustainability of specific land-use systems, we need approaches for the effective assessment of the present species richness and models that can predict the effects on species richness as realistically as possible. In this context, we present a methodology to estimate and predict vascular plant species richness at the local and the regional scale. In our approach, the major determinants of vascular plant species richness within the study area are taken into consideration: These are according to Duelli's mosaic concept the number of habitat types and of habitat patches within area units. Furthermore, it is based on the relative frequencies of species within habitat types. Our approach comprises six steps: (i) the determination of present habitat patterns within an observation area, (ii) the creation of a land-use scenario with simulated habitat patterns, (iii) the determination of species frequencies within habitat types of this area, (iv) a grouping of habitat-specific species, (v) the estimation of the probabilities for all species (or habitat specialists) to occur, either in stepwise, exponentially enlarged landscape tracts (local scale), or in the entire observation area (regional scale), and (vi) the validation of the estimated species numbers. The approach will be exemplified using data from the municipal district of Erda, Lahn-Dill Highlands, Germany. The current species numbers to be expected on the basis of probability calculations were compared with those recorded on the basis of extensive field work. This comparison shows that, on the basis of our simple calculations, the current local plant species richness can be predicted well, with a slight underestimation. This revised version was published online in May 2005 with corrections to the Cover Date. This revised version was published online in July 2006 with corrections to the Cover Date.     

黄土高原旱地苹果园生草对土壤水分的影响

 对黄土高原渭北地区旱作生草苹果园3～10月0～100 cm土层土壤水分观测结果表明: 在0～40 cm土层牧草与果树存在水分竞争, 但生草对40～80 cm土层水分具有调蓄作用, 生草种类不同, 对土壤贮水增减量的影响存在差异, 其影响主要发生在春季与秋季, 在降水丰水年影响较小, 但在降水欠水年生草对果园土壤贮水影响较大。     

Association of vegetation and soil mite assemblages with isolated Scots pine trees on a Scottish wet heath

Isolated trees may significantly enhance biodiversity at the landscape level. However, our understanding of their impacts is still poor, particularly in environments with high soil moisture where research on this topic has been comparatively limited. We examined understorey vegetation and soil oribatid mite assemblages under live and dead Scots pine trees and in open treeless areas, all within the same Scottish upland wet heath system, to determine whether isolated live trees affected the understorey and mite components of the ecosystem, and whether these effects occurred in parallel. We also explored whether these responses might result from tree-driven reductions in soil moisture content. Live trees reduced soil moisture (relative to wet heath and beneath dead trees) and appeared to change vegetation from wet heath to dry heath type communities. These effects were strongly related to tree trunk diameter (tree size). No major effects of dead trees on understorey vegetation or soil moisture were apparent. Higher mite species abundance and richness were found under live trees than in treeless open heath. Although mite abundances were lower under dead trees than live trees, richness remained similar, thus different factors seem to be regulating mite abundance and community composition. These findings indicate that landscape-level biodiversity responses to environmental change such as habitat fragmentation cannot be predicted from vegetation patterns alone, and that even in heavily fragmented landscapes comparatively small patches such as isolated individual trees can enhance biodiversity.     

Evidence of Hierarchical Patch Dynamics in an East African savanna?

The Hierarchical Patch Dynamics Paradigm provides a conceptual framework for linking pattern, process and scale in ecosystems, but there have been few attempts to test this theory because most ecological studies focus on only one spatial scale, or are limited in their temporal scope. Here I use palaeoecological techniques (analysis of fossil pollen and stable carbon isotopes) to compare vegetation heterogeneity in an east African savanna at three spatial scales, over hundreds of years. The data show that patterns of vegetation change are different at the three spatial scales of observation, and suggest that different ecological processes dominate tree abundance at micro, local and landscape scales. Interactions between plants, disturbance (e.g., by fire and herbivores), climate and soil type may influence tree density at differing spatial and temporal scales. This hierarchical explanation of savanna vegetation dynamics could inform future biodiversity conservation and management in savannas. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evidence of Hierarchical Patch Dynamics in an East African savanna?

The Hierarchical Patch Dynamics Paradigm provides a conceptual framework for linking pattern, process and scale in ecosystems, but there have been few attempts to test this theory because most ecological studies focus on only one spatial scale, or are limited in their temporal scope. Here I use palaeoecological techniques (analysis of fossil pollen and stable carbon isotopes) to compare vegetation heterogeneity in an east African savanna at three spatial scales, over hundreds of years. The data show that patterns of vegetation change are different at the three spatial scales of observation, and suggest that different ecological processes dominate tree abundance at micro, local and landscape scales. Interactions between plants, disturbance (e.g., by fire and herbivores), climate and soil type may influence tree density at differing spatial and temporal scales. This hierarchical explanation of savanna vegetation dynamics could inform future biodiversity conservation and management in savannas. This revised version was published online in July 2006 with corrections to the Cover Date.     

Simulated productivity of heterogeneous patches in Southern African savanna landscapes using a canopy productivity model

A daily model of terrestrial productivity is used to simulate the annual productivity of heterogeneous vegetation structure at three savanna/woodland sites along a large moisture gradient in southern Africa. The horizontal distributions of vegetation structural parameters are derived from the three-dimensional canopy structure generated from detailed field observations of the vegetation at each site. Rainfall and daily climatic data are used to drive the model, resulting in a spatially explicit estimate of vegetation productivity in 100 m² patches over an area 810,000 m² (8,100 patches per site). Production is resolved into tree and grass components for each subplot. The model simulates the relative contribution of trees and grasses to net primary productivity (NPP) along the rainfall gradient. These simulated production estimates agree with previously published estimates of productivity in southern African savannas. Water-use efficiency of each site is directly related to the structural composition of the site and the differing water-use efficiencies for tree and grass functional types. To assess the role of spatial scale in governing estimates of vegetation productivity in heterogeneous landscapes, spatial aggregation is performed on the canopy mosaic at the northern-most (wettest) site for 625 m², 2500 m² and 5625 m² resolutions. These simulations result in similar overall patterns of average NPP for both trees and grasses, but drastically reduced distributions of productivity due to reduced structural heterogeneity. In particular, the aggregation of the detailed spatial mosaic to coarser resolutions is seen to eliminate information regarding demographic processes such as regeneration and mortality, and the dependence of grass productivity on over-story density. These results indicate that models of system productivity in savanna/woodland ecosystems must retain high spatial resolution to adequately characterize multi-year structural responses and to accurately represent the contribution of grass biomass to overall ecosystem production.This revised version was published online in May 2005 with corrections to the Cover Date.     

Toward a rule-based biome model

Current projections of the response of the biosphere to global climatic change indicate as much as 50% to 90% spatial displacement of extratropical biomes. The mechanism of spatial shift could be dominated by either 1) competitive displacement of northern biomes by southern biomes, or 2) drought-induced dieback of areas susceptible to change. The current suite of global biosphere models cannot distinguish between these two processes, thus determining the need for a mechanistically based biome model. The first steps have been taken towards the development of a rule-based, mechanistic model of regional biomes at a continental scale. The computer model is based on a suite of empirically generated conceptual models of biome distribution. With a few exceptions the conceptual models are based on the regional water balance and the potential supply of water to vegetation from two different soil layers, surface for grasses and deep for woody vegetation. The seasonality of precipitation largely determines the amount and timing of recharge of each of these soil layers and thus, the potential mixture of vegetative life-forms that could be supported under a specific climate. The current configuration of rules accounts for the potential natural vegetation at about 94% of 1211 climate stations over the conterminous U.S. Increased temperatures, due to global warming, would 1) reduce the supply of soil moisture over much of the U.S. by reducing the volume of snow and increasing winter runoff, and 2) increase the potential evapotranspiration (PET). These processes combined would likely produce widespread drought-induced dieback in the nation’s biomes. The model is in an early stage of development and will require several enhancements, including explicit simulation of PET, extension to boreal and tropical biomes, a shift from steady-state to transient dynamics, and validation on other continents.     

不同土壤水分条件下草莓果实在模拟贮运中的损伤易感性

通过模拟运输振动实验研究了种植在不同土壤水分条件下草莓果实在贮运中的损伤易感性。试验结果表明,栽培在较高土壤水分条件下的果实含有较高的水分,贮运中果实损伤易感性与果实含水量、果实鲜重、土壤湿度呈显著或极显著相关。模拟振动实验发现:在高土壤水分条件下栽培的草莓果实在振动前、振动后及贮藏后的呼吸速率分别为生长在低土壤水分果实的124.9%、119.0%和133.2%。振动的果实比未振动的果实降低了28%-36%的蔗糖,增加了4.5%-9.8%的果糖和7.1%-10.1%葡萄糖,说明运输振动促进了蔗糖降解为葡萄糖和果糖的水解过程。     

蔗叶不同还田模式对土壤水分和甘蔗产量的影响

田间试验研究了蔗叶不同还田模式对旱地蔗园土壤水分和甘蔗产量的影响。结果表明,蔗叶不同还田模式的甘蔗产量差异不显著,蔗叶全覆盖还田与蔗叶隔行覆盖还田处理的总苗数较焚烧处理差异达极显著性水平（P〈0.01）,有效茎与焚烧处理差异达显著性水平（P〈0.05）。蔗叶全覆盖还田与蔗叶隔行覆盖还田对宿根蔗垄上前期与后期的土壤水分保持效果较好,不同蔗叶还田模式对甘蔗行间的土壤水分保持效果较好。