Carex sempervirens tussocks induce spatial heterogeneity in litter decomposition, but not in soil properties, in a subalpine grassland in the Central Alps

Tussocks of graminoids can induce spatial heterogeneity in soil properties in dry areas with discontinuous vegetation cover, but little is known about the situation in areas with continuous vegetation and no study has tested whether tussocks can induce spatial heterogeneity in litter decomposition. In a subalpine grassland in the Central Alps where vegetation cover is continuous, we measured soil properties [concentration of N, C, organic matter (OM) and pH] and monitored litter decomposition traits (dry mass loss, loss of C, N, P and K) inside and outside tussocks of Carex sempervirens. Soil C, N, OM concentrations or pH inside tussocks did not differ significantly from those outside tussocks. After 1 year of decomposition, litter dry mass loss, C and K loss were significantly smaller inside than outside tussocks. The slower litter decomposition inside tussocks was likely caused by the elevated tussock base, which made environmental conditions inside tussocks much dryer than those outside in early spring when snow melts. Our results suggest that in areas with continuous vegetation cover tussocks induce spatial heterogeneity in litter decomposition but not in soil properties.     

草地物种多样性对群落可入侵性影响的研究进展

综述草地群落入侵实验中物种多样性和群落可入侵性关系的研究进展。目前物种多样性与群落可入侵性主要出现了对立的关系模式,被普遍接受的解释机制为尺度依赖。但其它研究中出现了更为复杂的关系,提出物种特性、植物更新、种间关系变化和群落构建机制等其它因素可能是导致物种多样性与群落可入侵性出现复杂关系的原因。建议未来研究中应注意的几个问题,即物种多样性与群落可入侵性关系在不同营养级适用性,与群落构建机制变化间的联系和时间尺度对物种多样性与群落可入侵性关系的影响。     

Relationship between group dynamics and spatial distribution of African elephants in a semi‐arid environment

Group dynamics related to distribution of African elephants in the semi‐arid environment of Tsavo East National Park, Kenya was studied between 2007 and 2010. I examined the seasonal distribution of lone bulls, bull groups, family units and mixed groups and group size. Lone bulls were widely distributed in the dry season and localized in the wet season, whereas bull groups were localized in both seasons. On average bull group size did not differ with seasons or areas, but larger groups were observed in preferred areas in the dry season. Family units were widely distributed in the dry season and localized in the wet season, whereas mixed groups were localized in both seasons. Although family units and mixed groups tended to be large in all areas in the wet season, large groups were also observed in the dry season. Large family units were widely distributed, whereas large mixed groups were localized. The relationship between group dynamics and distribution was attributed to the combination of social, ecological and poaching factors. This study provided insight into distribution patterns that can be applied to the security of the Tsavo elephant population.     

Determining adrenocortical activity as a measure of stress in African elephants (Loxodonta africana) in relation to human activities in Serengeti ecosystem

African elephants (Loxodonta africana) play a vital role in most African ecosystems, with their opportunity to alter the entire ecosystem by their sheer numbers. Defining and measuring animal welfare has been much discussed. One potential way of determining an animal's welfare is to record the absence or presence of stress. Little research on elephant welfare has so far been performed in the Serengeti ecosystem. The aim of this study was to record the faecal glucocorticoid metabolite levels of African elephants in areas with high or with minimum human interference. A total of 117 faecal samples were collected from randomly located single elephants as well as family herds in the northern, central and western Serengeti National Park (SNP) as well as in Grumeti Game Reserve and Ikoma Open Area, northern Tanzania in 2010. Elephants had higher levels of faecal glucocorticoid metabolites in the areas outside, compared with areas inside SNP. No single males were observed outside SNP, and in general, higher abundance of elephants was observed inside SNP. This suggests that elephants may prefer to reside in the potential safer areas inside the national park, demonstrating the importance of protected areas to improve the welfare of elephants.     

Effects of wind erosion on the spatial heterogeneity of soil nutrients in two desert grassland communities

Wind is known to affect the spatial heterogeneity of soil resources in arid and semiarid systems, but multi-year, quantified observations are largely absent. We studied the effects of wind erosion on the spatial distribution of soil organic carbon (SOC) and other soil nutrients at the Jornada Experimental Range, in southern New Mexico. Enhanced wind erosion was encouraged by grass cover reduction in a Sporobolus-mesquite dominated site (SM) and a Bouteloua-mesquite dominated site (BM). The scale and magnitude of spatial dependence for the soil analytes were quantified using geostatistical analyses. Results of this study show that soil organic matter related analytes such as SOC, TN, N_avail, and SO₄ ^2- are among the first to be eroded and redistributed; cations such as Ca²⁺ and Mg²⁺ may not be removed and redistributed significantly; and other ions such as K⁺, Na⁺ and Cl⁻ showed no discernible pattern of change. Geostatistics show that wind appeared to increase the scale of spatial autocorrelation, but decrease the scale of spatial dependence of most soil analytes over 2–3 windy seasons. In the wind enhanced plot of the SM site, up to 99% of the spatial dependence of SOC was autocorrelated at the distance of 1.45 m before the initiation of wind erosion, but the spatial dependence dropped significantly to only 60% at a larger autocorrelation distance of 2.76 m after three windy seasons. Similar but less significant changes were observed for SOC in the BM site. Despite the differential effects of wind on the soil analytes, we conclude that the overall results of wind on the grass cover reduction plots are the disappearance of small, strong fertile islands, which may be related to grasses; and the reinforcement of large fertile islands, which are likely related to mesquite shrubs. In addition, the change of the spatial patterns of SOC and other soil nutrients induced by enhanced wind erosion may persist and reinforce soil islands associated with shrubs, thus allowing a positive feedback for further desertification in this arid grassland.     

桂西北喀斯特常绿落叶阔叶混交林物种多样性分布格局的尺度效应

物种多样性的空间分布格局及其与尺度的关系研究对于了解群落物种多样性形成机制具有重要意义。为了探讨喀斯特地区物种多样性空间分布格局的尺度效应,以喀斯特常绿落叶阔叶混交林25 hm~2样地的(胸径DBH≥1)木本植物为研究对象,分析了6个空间尺度(5 m×5 m,10 m×10 m,20 m×20 m,50 m×50 m,100 m×100 m,250 m×250 m)上的多度、物种丰富度、Shannon-Wiener指数、Simpson指数以及Pielou均匀度指数的变化规律。结果表明:物种多样性指数的空间分布均表现出较高的空间异质性;物种多样性指数的方差随取样尺度增加呈现单峰分布特征,并且在100 m×100 m尺度上达到最大值;物种多样性指数的变异系数随尺度的增加呈线性下降趋势,其中,Shannon-Wiener指数、Simpson指数以及Pielou均匀度指数均在5 m×5 m至20 m×20 m尺度上明显减小;在大于50 m×50 m的尺度上,物种丰富度与多度的正相关性不显著(P0.05)。喀斯特常绿落叶阔叶混交林物种多样性的空间分布格局与不同空间尺度密切相关,深入解析物种多样性随空间尺度的变化模式,需要在类似的森林生态系统做更多的研究。     

Underlying effects of spatial aggregation (clumping) in relationships between plant diversity and resource uptake

In the debate on the influence of declining plant diversity on ecosystem functioning, spatial aggregation has received little attention, in spite of the fact that the local distribution of species determines interaction. Using a simple conceptual model, we visualize the effect that clumping has on below ground resource uptake (RU). Two general conclusions can be drawn from the model: (1) clumping results in a loss of RU, and does so to a larger extent when species differ more in rooting depth and when plant density increases; (2) the current view on complementarity may be flawed, as we find that neighborhood species richness only promotes spatial complementarity directly at the transition from monocultures to higher species richness levels, but not at transitions between higher richness levels. The number of species in a community does, however, affect the probability of the occurrence of clumping. In randomly assembled communities, clumps are statistically larger when the number of species is lower, leading to reduced overall below ground complementarity. Complementarity, and thus also RU, will therefore increase indirectly with increasing species richness. As a result of our findings, we propose that clumping could have a number of 'hidden' consequences, for example on invasibility of a community and on competition between species. Clumping could also prove to be a new and important explanatory factor in diversity studies, as it extends the definition of complementarity effects to include the effect of spatial heterogeneity of plant positions within the community.     

Seasonality regulates the effects of resource addition on plant diversity and ecosystem functioning in semi-arid grassland

季节性调控资源添加对半干旱草原物种多样性与生态系统功能的影响在资源富集的条件下,物种丰富度、群落地上生产力以及群落稳定性的季节变化通常被忽视。本研究致力于探究在干旱区草原,资源添加如何在生长季的不同月份影响物种丰富度、群落地上生产力及其稳定性。我们在内蒙古草原设置了为期3年的资源添加(氮添加-N、水添加-W以及水、氮共同添加-NW)实验,利用季节性取样的方法,去检验资源添加(水、 氮)对物种丰富度、群落地上生产力及其稳定性的影响,并通过构建结构方程模型分析资源添加调控不同月份与整个生长季群落稳定性的内在机制及其相对重要性。研究结果表明,资源添加在整体上未改变5与6月的群落地上生产力,而氮与水氮共同添加显著提高了7与8月的群落地上生产力。资源添加在整体上未改变物种丰富度、物种异步性与群落稳定性。氮添加与水氮共同添加提高了7与8月的群落地上生产力,主要源于其增加了多年生丛生禾草的地上生产力。结构方程模型分析表明：在生长季前期与整个生长季,物种异步性是决定群落稳定性的主要机制;在生长季后期,多年生丛生禾草的稳定性是驱动群落稳定性的主要因子。我们的研究证明：在半干旱草原,季节与资源的有效性可以交互影响群落地上生产力及其稳定性。这些发现对于半干旱草原的季节性可持续管理具有重要意义,以期减轻土地利用与全球变化带来的影响。     

On the scaling patterns of species spatial distribution and association

Understanding how species distribution (occupancy and spatial autocorrelation) and association (that is, multi-species co-distribution) change across scales is fundamental to unlocking the pattern formation in population ecology and macroecology. Based on the Bayesian rule and join-count statistics, I present here a mathematical model that can demonstrate the effect of spatial scale on the observation of species distribution and association. Results showed that the intensity of spatial autocorrelation and species association declines when the grain in the spatial analysis increases, although the category of species distribution (aggregated or segregated) and association (positive or negative) remains the same. Random distribution and species independence were proved to be scale-free. Regardless of the possible patterns of species distribution and association, species tend to be randomly distributed and independent from each other when scaling-up (an increasing grain), reflecting a percolation process. This model, thus, grasps the statistical essence of species scaling pattern and presents a step forward for unveiling mechanisms behind species distributional and macroecological patterns.     

海南尖峰岭热带山地雨林物种多样性空间分布格局的尺度效应

物种多样性的空间分布格局和维持机制是群落生态学的基本问题。为了探讨海南尖峰岭地区物种多样性空间分布格局的尺度效应, 以海南尖峰岭热带山地雨林60 hm²样地为研究对象, 分析了物种丰富度、物种多度、Shannon-Wiener指数、Simpson指数以及Pielou均匀度指数随6个空间取样尺度(5 m × 5 m、10 m × 10 m、20 m × 20 m、40 m × 40 m、100 m ×100 m、200 m × 200 m)的变化。结果表明: 相比Simpson指数和Pielou均匀度指数, 物种丰富度、多度以及Shannon-Wiener指数具有更为明显的空间尺度效应; 物种丰富度的方差随取样尺度增加呈现单峰分布特征, 并且在20 m × 20 m尺度上达到最大值, 而物种多度的方差随取样尺度的增加而增大; 物种丰富度和多度的正相关性随着取样尺度的增加逐渐减小甚至消失, 这可能与随取样尺度增加生境异质性增加有关; 取样尺度对3个物种多样性指数空间分布的影响可能与研究区域内稀有种的组成有关。     

Topographic and ungulate regulation of soil C turnover in a temperate grassland ecosystem

A major obstacle for predicting the effects of climate and land use changes on global soil carbon (C) stores is the very limited knowledge about the long timescale dynamics of the relatively stable fraction of soil C, which represents the bulk of soil C and the primary determinant of the long‐term C balance of terrestrial ecosystems. In this study, we examined how variable topo‐edaphic conditions and herds of native migratory ungulates influenced turnover of the stable pool (total minus active fraction) of soil C in grasslands of Yellowstone National Park (YNP). Soil C properties were determined for grasslands located inside and outside long‐term ungulate exclosures established 1958–1962 at seven variable topographic positions. Active C pool sizes, estimated with soil laboratory incubations, and soil radiocarbon measures were used to parameterize a process‐based model to determine turnover of the stable C pool at the sites. Stable C turnover ranged 37–653 and 89–869 years for 0–10 and 0–20 cm soils, respectively. Among ungrazed communities, there was a trend for stable soil C turnover to slow along topographic gradients of increasing soil moisture, soil C content, and shoot biomass from hilltop to slope‐bottom positions. This was likely a result of an increasing amount of support tissue resulting in greater concentrations of lignin and cellulose as shoot biomass increased down slope. In contrast, across the grazed landscape, stable C turnover sped up from hilltop to slope‐bottom positions, which was likely a consequence of grazer effects on plant species composition along the topographic gradient. These findings indicated that despite topography playing the primary role in controlling such important site characteristics as soil moisture, soil C content, and plant production in YNP grassland, the long‐term turnover of the stable C pool was determined by herbivores. The results demonstrate the important regulatory role of herbivores in controlling the C balance of this semiarid grassland ecosystem.     

Relationship between spatial strategies and morphological attributes in a Uruguayan grassland: a functional approach

Abstract. In a native grassland in Uruguay subjected to continuous grazing by cattle, four permanent plots of 40 × 40 cm were established. Plots were divided into 16 × 16 cells and the presence of species was recorded seasonally during two years. The spatial dynamic of the dominant species was defined through three parameters: persistence, short-distance spread and long-distance spread. The association between the spatial strategies and certain morphological attributes that were presumed to be important to the spatial behaviour of the species, was examined using Correspondence Analysis. Four types of spatial strategies were observed: (1) persisting over the year in the same cell without showing any type of mobility; (2) continuously growing due to the solid advancing front of ramets, persisting or not persisting (3) in the same cells; and (4) multi-strategy with high persistence and both high short- and long-distance spread. Among the selected attributes, the erect/prostrate habit, the capacity of vegetative propagation by means of bulbs, rhizomes or stolons, the ratio length/width of leaves and the season of active growth were the characteristics most closely associated with the spatial strategies.     

Below-ground spatial pattern of rhizomes in a grassland community and its relevance to above-ground spatial pattern

Studies of spatial patterns in grassland plant communities have focused on above-ground patterns, ignoring the fact that in clonal plant communities, such as those found in grasslands, above-ground spatial structure must reflect below-ground horizontal growth. The present study examines, at both a fine and a coarse spatial scale, relationships between rhizome and ramet distribution. At the coarse scale, the dominance of species differed between above- and below-ground; some species dominated only above- or below-ground, and others dominated in both layers. At the fine scale, a single species' ramet aggregation above-ground significantly differed from its rhizome aggregation below-ground, for many species. Even for a given species, quantitative relationships between above- and below-ground dominance varied among localities. The variation in spatial pattern among species can be explained by attributes of clonal growth form, including rhizome size, rhizome origin and pattern of above-ground ramet aggregation. Using these parameters of clonal growth, four major space occupation patterns were described for mountain grassland species. For species with a high abundance of evenly distributed rhizomes, ramets either i) reflect rhizome distribution, or ii) do not fully reflect rhizome distribution, but rather are spatially aggregated, and rhizomes are typically developed below-ground. For species with a low abundance of rhizomes, ramets either iii) reflect rhizome distribution and rhizomes are typically of above-ground origin, or iv) do not reflect rhizome distribution and are aggregated only at the growing tips of rhizomes. Spatial correlation above-ground among species was the same as below-ground for some pairs of species but was significantly different for other pairs. This revised version was published online in June 2006 with corrections to the Cover Date.     

Optimal movement between patches under incomplete information about the spatial distribution of food items

If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed and therefore always moves to the nearest patch (be it empty or filled). In contrast, a prescient forager is fully informed and only exploits filled patches, skipping all empty patches. A Bayesian assessor has prior knowledge about the content of patches (i.e. it knows the characteristics of the spatial pattern) and may skip neighbouring patches accordingly by moving to the patch where the highest gain rate is expected. In most clumped and regular distributions there is a benefit of assessment, i.e. Bayesian assessors achieve substantially higher long-term gain rates than ignorant foragers. However, this is not the case in distributions with less strong spatial pattern, despite the fact that there is a large potential benefit from a sophisticated movement rule (i.e. a large penalty of ignorance). Bayesian assessors do also not achieve substantially higher gain rates in environments that are relatively rich or poor in food. These results underline that an incompletely informed forager that is sensitive to spatial pattern should not always respond to existing pattern. Furthermore, we show that an assessing forager can enhance its long-term gain rate in highly clumped and some specific near-regular food distributions, by sampling the environment in slightly larger spatial units.     

Impact of cattle dung deposition on the distribution pattern of plant species in an alvar limestone grassland

Abstract. Seed banks in cattle dung, soil under cattle dung and soil under vegetation and growth response of plant species to the changes in soil nitrogen availability were studied in an alvar limestone grassland on Öland, Sweden, in order to analyse the impact of dung deposition and decomposition on the formation of patches of plant species. Results suggest that patches of four plant species could result from cattle dung deposition and decomposition. Impact of dung could proceed in three ways: (1) by changing the relative abundance of species in the soil seed bank under dung, and/or (2) by influencing the deposition of seeds in the dung, and/or (3) by intensifying the growth of some species through nutrient release. Species patches could result from one or more of these aspects. For instance, patches of Arenaria serpyllifolia may be induced by dung deposition because of the dominance of its seeds in dung, while the pattern of Cerastium semidecandrum and Festuca ovina may be due to the abundance of their seeds in the soil seed bank under dung and their positive growth response to increased nitrogen availability.     

Landscape heterogeneity metrics as indicators of bird diversity: Determining the optimal spatial scales in different landscapes

Species distribution models are often used to study the biodiversity of ecosystems. The modelling process uses a number of parameters to predict others, such as the occurrence of determinate species, population size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial scales when it comes to undertaking a GIS analysis.We explored the goodness of fit of several models using the metrics of landscape heterogeneity and altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a breeding bird survey by means of point counts. The relationships between biodiversity and landscape metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 different spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial scale for each landscape typology.Our results revealed that the relationships between species’ richness and landscape heterogeneity using 1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of landscape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas. The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over extensive areas.