Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

干旱半干旱区灌丛斑块与降水量响应关系的熵模型模拟

灌丛斑块分布格局是灌木在干旱缺水条件下对生存环境的自我调节和适应的具体表现。应用熵理论和Klausmier模型,解释了灌丛斑块水分聚集原理并模拟了不同年降水条件下灌丛斑块的最佳面积比值(即最佳灌丛盖度)。研究结果表明:灌丛斑块生物量与其土壤含水量呈反比例函数关系,当生态系统处于稳定状态时(即熵最大状况下),年降水量与灌丛斑块面积比值符合一定的线性关系。研究采用内蒙古草原地区的野外调查数据,获得模型所需参数,进而模拟了不同年降水量条件下灌丛斑块最佳面积比值,研究结果可为半干旱地区植被保护与恢复提供参考。     

Ecosystem Management of Desertified Shrublands in Israel

The objectives of this study were to understand the ecological processes and possible management strategies in desertified shrublands. We hypothesized that biological production and diversity in desertified shrublands in the Negev in Israel are low due to water, soil, and nutrient leakage from the ecosystem. We designed a series of field experiments in order to examine (a) whether source–sink relationships exist between the crusted soil and the shrub patches, (b) whether resources (water, soil, and nutrients) leak from the system, and (c) whether management, which changes the landscape mosaic by introducing new sink patches that reduce leakage of resources, may increase productivity and diversity. The results indicate that the low number of shrub patches, which serve as sinks for resources, leads to water, soil, and nutrient leakage from the ecosystem. This leakage reduces ecosystem production and diversity. We found that artificially created pits, which act as sinks for resources, decrease leakage and increase biomass production and annual plant species diversity. Based on the experimental results, we developed conceptual models for shrubland desertification and ecosystem management. The models are based on a source–sink relationship between two patch types characteristic of shrublands. The models relate landscape productivity to the number of sink patches and suggest that, in cases where there are too few sinks, artificially created sink patches should be added. Management methods were developed to reduce resource leakage in the desertified shrubland of the Negev. Methods included construction of man-made pits in the landscape that add resource-enriched patches to the landscape. These patches are used to create parks consisting of clusters of trees integrated into a matrix of shrubs and herbaceous vegetation. The managed parks are used for recreational purposes and for rangeland. Received 8 July 1997; accepted 7 July 1998.     

Grassland connectivity explains entomophilous plant species assemblages in an agricultural landscape of the Pampa Region,Argentina

The Pampa grassland of Argentina is one of the most highly threatened biomes in the world. A high proportion of the original grassland cover has been transformed into land for agriculture or degraded. In the southern part of the region, fragmented semi‐natural grasslands over exposed rock still persist and connectivity between them is assumed to be crucial for maintaining viable populations. We quantified overall connectivity of grassland patches in a sector of the Southern Pampa region, and investigated the degree to which landscape connectivity explains entomophilous plant species assemblages in a subset of patches. We characterized each of the 301 patches in the landscape by their degree of intra‐patch and inter‐patch connectivity based on graph theory, and considering threshold dispersal distances from 100 to 1000 m. We surveyed entomophilous plant species in 39 grassland patches and classified the species in three categories (annual herbs, perennial herbs and shrubs) considering their different growth form and longevity. The influence of connectivity variables on entomophilous plant species assemblages variation was explored using Canonical Correspondence Analysis. Although grassland patches were poorly connected at all threshold distances, some of them were found to be critical for global connectivity. Connectivity significantly explained total, annual‐biennial and shrub assemblages for all threshold dispersal distances (6–13% of total variation). Variation in annual species assemblages was associated with intra‐patch and inter‐patch connectivity at short distance (100 m), while variation in shrub species assemblages was explained by intra‐patch and inter‐patch connectivity for distances between 100 m and 1000 m. This study evidenced the low connectivity of the study system, allowed the identification of critical areas for conservation, and provided valuable information to develop management strategies in increasingly human‐dominated landscapes.     

Role of Salix reinii patches in spatio-temporal patterns of cohabitants on a Japanese volcano

深山柳斑块在日本火山共栖物种时空格局中的作用 灌木斑块通常能为定植物种提供支持，尤其是在具有压力和/或干扰的环境中，这种作用更明显。山地上的压力和干扰强度通常与海拔梯度有关，并会因灌木斑块(树冠和凋落物)而降低。本研究旨在利用火山上不同海拔高度的灌木斑块来阐明这些关系。从2005年早春到2006年夏季，我们在日本北部高丽山(海拔1131 m)的3个海拔高度监测了深山柳(Salix reinii)斑块内外所有地块中的所有嫩枝及其相关 环境，例如温度、光照、湿度和化学成分。我们通过面积、分枝密度和凋落物厚度对斑块结构进行评价。研究结果表明，灌木斑块的结构在不同海拔之间并无差异。通过在整体上增加植物物种的丰富度，这些斑块提高了稀有物种的多样性。斑块内的水分、氮和磷含量均要高于斑块外。灌木斑块对于共栖物种的影响总体上是正向的，但对幼苗丰度、嫩枝存活率和开花的影响则是负面的。这些结果表明灌木斑块的生长格局决定了植物群落结构，并促成了植物的多样性。     

Temporal asynchrony in fine‐root biomass may contribute to shrub and grass coexistence in mixed patches

We described seasonal changes in fine‐root biomass of a grass and a shrub dominant species in a plant community characteristic of the arid Patagonian Monte and then we inferred to want extent the observed differences could contribute to the species coexistence. We selected representative plant patches of the natural vegetation arrangement consisting of one isolated plant of the dominant shrub Larrea divaricata (Ld), grass patches formed by one or more bunches of the dominant grass Nassella tenuis (Nt), and mixed patches consisting of one individual of L. divaricata with bunches of N. tenuis under its canopy (LdNt). We assessed the biomass and temporal changes in fine roots of each species in the upper soil (50 cm depth) of each patch type at three‐month intervals during 2 years. The temporal series of fine‐root biomass were compared among patch types and in relation to above‐ground phenology, as well as climate variables (precipitation, arid index and air temperature). Seasonal changes in fine‐root biomass showed similar cycles in the three plant patches with a maximum in spring. The maximum increase in root biomass in Ld and Nt patches occurred during the onset of reproductive growth in winter and spring, respectively. Fine‐root changes in LdNt patches mimicked that in Ld patches. Precipitation inputs were significantly positively and temperature negatively related to fine‐root changes in Nt patches. Fine‐root changes in Ld and LdNt patches were related to the aridity index (positively) and temperature (negatively). We concluded that the observed asynchronies in the date of the largest increases in root biomass and its climate control between the studied grass and shrub species could contribute to the coexistence of plants of both life forms when they overlap their root systems growing in mixed patches. Mechanisms underlying the root patterns observed should be further explored.     

Deficit in community species richness as explained by area and isolation of sites

Abstract . The potential community species richness was predicted for 85 patches of seminatural grassland in an agricultural landscape in Denmark. The basis of the prediction was a very large dataset on the vegetation, soil pH and topography in Danish grasslands and related communities. Species were inserted into potential species pools according to their preferences regarding soil acidity and water availability (expressed as potential solar irradiation), and to the ranges in these two factors observed in each grassland patch. The difference between the predicted and the observed patch‐level species richness, community richness deficit, varied considerably among patches. Community richness deficit exhibited a negative relationship with patch area, and for small patches a positive relationship with patch isolation.     

三江平原残存湿地斑块特征及其对物种多样性的影响

景观破碎化是当前一个突出的生态学问题,破碎斑块的大小和形状特征对物种多样性具有重要影响。选取三江平原别拉洪河流域为研究区,通过调查残存湿地斑块的面积、形状和植物物种丰富度,利用相关分析和回归分析等方法研究了残存湿地斑块特征及其对物种多样性的影响。研究结果表明:在强烈的人为活动干扰下,三江平原湿地破碎化严重,残存湿地斑块面积较小,斑块形状规则、边界简单。斑块形状指数、分维数、形状特征点数和斑块面积、物种丰富度均没有显著相关关系,而斑块面积、周长和周长面积比对物种丰富度具有显著影响。但是斑块周长和周长面积比均受面积的直接影响,所以斑块面积是决定物种丰富度的主要因素,这点需在湿地生物多样性保护中予以重视。虽然斑块形状特征点数是农业景观物种丰富度的一个很好预测指标,但是该指标不适用于形状规则的残存湿地斑块物种丰富度预测。另外,对于残存湿地斑块种-面积关系的机理还有待进一步研究。     

Local and landscape‐scale effects of an ant nest construction in an open dry forest of Uruguay

1. Ants modify soil properties via nest construction and by doing this modulate soil resources for other organisms. In this sense ants are recognised as ecosystem engineers. 2. In this framework, two less well‐studied issues are focused on: (i) the permanence of the effects of ant nests on plant communities after colonies have died, and (ii) the scaling up from patch to landscape‐scale effects. 3. The aim of the present study was to address these issues in the open dry forests of Uruguay, inhabited by the ant Atta vollenweideri Forel. The active and abandoned nests of this ant represent different and conspicuous patches (30–60 m²) in the landscape. 4. The soil concentration of sodium, a key element in the system, was substantially higher among active nests, and remains high during the early stages of abandoned nests. Woody species abundance, richness, and composition were affected at the patch scale, and simulation models suggested an increase in species richness at the landscape scale. 5. The present study highlights the importance of abandoned nests for plant‐species richness in the ecosystem engineer framework and the need to advance in an integrative approach to study both local and landscape effects of ant's nests.     

Rethinking patch size and isolation effects: the habitat amount hypothesis

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non‐habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal‐sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal‐sized sample sites rather than for unequal‐sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch‐scale and landscape‐scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.     

Harvester ant response to spatial and temporal heterogeneity in seed availability: pattern in the process of granivory

The influence of temporal and spatial heterogeneity in seed availability on the foraging behaviour of the harvester ant Messor arenarius was studied in an arid shrubland in the Negev Desert, Israel. The study investigated the implications of behavioural responses to heterogeneity in seed availability for the seed predation process and the potential for feedback effects on vegetation. Vegetation and seed rain were monitored across two landscape patch types (shrub patches and inter-shrub patches) in 1997. Shrub patches were shown to have higher plant and seed-rain density than inter-shrub patches. Patch use and seed selection by M. arenarius foragers were monitored through the spring, summer and autumn of 1997. After a pulse of seed production in the spring, the ants exhibited very narrow diet breadth, specialising on a single annual grass species, Stipa capensis. At this time, ants were foraging and collecting seeds mainly from inter-shrub patches. In the summer, diet breadth broadened and use of shrub patches increased, although the rate of seed collection per unit area was approximately equal in the two patch types. The increase in the use of shrub patches was due to colony-level selection of foraging areas with relatively high shrub cover and an increase in the use of shrub patches by individual foragers. In the autumn, a pulse of seed production by the shrub species Atractylis serratuloides and Noaea mucronata led to a reduction in diet breadth as foragers specialised on these species. During this period, foragers exhibited a large increase in the proportion of time spent in shrub patches and in the proportion of food items collected from shrub patches. The seasonal patterns in foraging behaviour showed linked changes in seed selection and patch use resulting in important differences in the seed predation process between the two landscape patch types. For much of the study period, there was higher seed predation pressure on the inter-shrub patches, which were of relatively low productivity compared with the shrub patches. This suggests that the seed predation process may help maintain the spatial heterogeneity in the density of ephemeral plants in the landscape.     

Landscape‐ and small‐scale determinants of grassland species diversity: direct and indirect influences

Species richness is influenced both by mechanisms occurring at landscape scales, such as habitat availability, and local‐scale processes, that are related to abiotic conditions and plant–plant interactions. However, it is rarely tested to what extent local species richness can be explained by the combined effect of factors measured at multiple spatial scales. In this study, we quantified the simultaneous influence of historical landscape‐scale factors (past human population density, and past habitat availability – an index combining area and connectivity) and small‐scale environmental conditions (shrub cover, and heterogeneity of light, soil depth, and other soil environmental variables) on plant species richness in dry calcareous grasslands (alvars). By applying structural equation modelling (SEM) we found that both landscape conditions and local environmental factors had significant direct and indirect (i.e. through the modification of another factor), effects on species richness. At the landscape scale, we found a direct positive influence of historical habitat availability on species richness, and indirect positive influence of past human population (via its effects on historical habitat availability). At small scales, we found a positive direct influence of light heterogeneity and shrub cover on species richness. Conversely, we found that small‐scale soil environmental heterogeneity, which was mainly determined by soil depth heterogeneity, had a negative effect on species richness. Our study indicates that patterns of species richness in alvar grasslands are positively influenced by the anthropogenic management regime that maintained the landscape habitat conditions in the past. However, the abandonment of management, leading to shrub invasion and increased competition for light resources also influenced species richness. In contrast to the positive heterogeneity–diversity relationship we found that soil heterogeneity reduced species richness. Environmental heterogeneity, occurring at the plant neighbourhood scale (i.e. centimetres), can increase the isolation among suitable soil patches and thus hinder the normal functioning of populations. The combination of previous knowledge of the system with new ecological theories facilitates disentangling how species richness responds to complex relationships among factors operating at multiple scales.     

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.     

Influence of light and soil moisture on Sierran mixed-conifer understory communities

Sierra Nevada forests have high understory species richness yet we do not know which site factors influence herb and shrub distribution or abundance. We examined the understory of an old-growth mixed-conifer Sierran forest and its distribution in relation to microsite conditions. The forest has high species richness (98 species sampled), most of which are herbs with sparse cover and relatively equal abundance. Shrub cover is highly concentrated in discrete patches. Using overstory tree cover and microsite environmental conditions, four habitats were identified; tree cluster, partial canopy, gap, and rock/shallow soil. Herb and shrub species were strongly linked with habitats. Soil moisture, litter depth and diffuse light were the most significant environmental gradients influencing understory plant distribution. Herb cover was most strongly influenced by soil moisture. Shrub cover is associated with more diffuse light, less direct light, and sites with lower soil moisture. Herb richness is most affected by conditions which influence soil moisture. Richness is positively correlated with litter depth, and negatively correlated with direct light and shrub cover. Disturbance or management practices which change forest floor conditions, shallow soil moisture and direct light are likely to have the strongest effect on Sierran understory abundance and richness.     

Adjacency arrangement effects on plant diversity and composition in woodland patches

To understand "adjacency arrangement" (configuration of a patch and its adjoining elements) as a basic unit of landscape pattern, we studied 30 woodland patches ( Populus tremuloides) in Alberta, Canada, with adjacent vegetation varying from 0% shrubland (100% grassland) to 100% shrubland (0% grassland). We evaluated (1) how important adjacency effects are relative to resource availability and disturbance in affecting plant species richness and composition in the woodland understory, and (2) which species and species groups change in abundance and presence with increasing percent shrubland in the adjacency arrangement. We found that, in addition to topography and previous grazing, adjacency arrangement is a major significant variable affecting the vegetation composition of woodland patches. Along a gradient from 0% to 100% adjacent shrubland, the species composition in woodland patches changed from an abundance of weedy and introduced plants to an abundance of moist-environment plants, the proportion of native species and perennials increased linearly, the number of "shrubland species" increased linearly, "grassland species" decreased linearly, and total species richness remained constant. Woodland vegetation did not differ significantly where adjacent shrubland was to the east or to the west of the wooded patch, nor between patches which did or did not experience a change in the adjacent shrubland in the recent past. We conclude that adjacency arrangement is an easily measured, highly promising concept for ecological understanding, as well as for land planning, design, conservation and management.     

The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.     

The effects of shrub patch sizes on the colonization of pioneer plants on the volcano Mount Koma,northern Japan

Vegetation recovery on Mount Koma, Hokkaido, Japan, has been slow after the catastrophic eruption in 1929, due to undeveloped soil and limited plant colonization. Nowadays, the seedling establishment is supported mostly by a nurse plant, Salix reinii forming shrub patches, facilitates the plant colonization. Although the effects of shrub patches should differ with patch sizes, the size effects have not been examined well. To examine the size effects, seed-sowing experiments were conducted on two common pioneer herbaceous species, Miscanthus sinensis and Polygonum sachalinense, in the field. The seed germination and seedling survival were monitored by the seeds sown into S. reinii patches (0.97 m²–4.12 m² in area) for 4 months during snow-free periods. Microenvironments altered by the patches were measured. Lab-experiments were performed to characterize the seed germination and seedling growth.Larger patches decreased light intensity and temperature more and increased litter and water content. The large patches promoted the seed germination of the two species. Interspecific interactions, examined by a seed mixture experiment, showed that the interaction increased the seed germination on M. sinensis and decreased that on P. sachalinense. On the lab-experiments at three temperatures (15, 20 and 25 °C), M. sinensis seeds germinated more at higher temperatures and obtained higher seedlings biomass. P. sachalinense germinated the seeds more at 20 °C and grew faster at lower temperatures. The total biomass of the two species was reduced by shade that intercepted 50% of light intensity. The seed germination and seedling growth of these two species became higher on litter with 2 cm in depth than on no litter. Soil water supported seed germination when the seeds of these two species were mixed while the water reduced the growth of P. sachalinense seedlings. Therefore, the dry soils were suitable for their growths. In all the treatments, P. sachalinense seedlings showed higher mortalities than M. sinensis.In conclusion, the large patches facilitated more to the colonization of pioneer plants via seed germination and growth. Large patches acted as a nursery supporting the natural regeneration in the disturbed area by improving litter accumulation, maintaining soil water, reducing strong light and/or protecting from heat.     

Interactions of the indigenous evergreen shrub Sabina vulgaris with coexisting species in the Mu Us sandland

Aims Facilitation is widespread in plant communities and particularly common in highly stressful environments. In the semi-arid Mu Us sandland where soil water and nutrients are short and grazing is heavy, many species grow within the large patches formed by the dense individuals of the unpalatable clonal shrub Sabina vulgaris, the only natural evergreen shrub species in the Mu Us sandland. However, the interactions between S. vulgaris and these coexisting species remain unclear, and we hypothesize that S. vulgaris can facilitate at least some species because within the patches vegetation is not grazed and soil water and nutrients may also be higher.Methods We measured the maximum height, coverage and number of individuals of each vascular species, little cover and thickness of biological crust at the soil surface in 1 m × 1 m plots in three types of microsites associated with 40 patches of S. vulgaris : (i) at the center, (ii) at the inner edge and (iii) outside the patches. We also took soil samples and measured soil water content and content of total N, P, K and organic matter.Important findings Soil water, nitrogen, organic matter and litter cover were the highest at the patch center, lowest outside the patches and intermediate at the inner edge of the patches, whereas thickness of biological crust was greater outside than at the center or at the inner edge. Among the 32 species recorded, six species preferred to occur within the patches, suggesting that S. vulgaris can facilitate these species most likely by grazing exclusion, increasing water and/or nitrogen resources in soil. However, most (19) species did not show preference and seven preferred outside. Also, species richness, pooled cover and number of individuals of all species were greater outside than within the patches, and such effects did not vary with the size of the S. vulgaris patches. These results suggest that the dominant interactions between S. vulgaris and the coexisting species are competition. The findings add to our knowledge that facilitation can be shown even when the competitive effects from the potential nurse plants are very strong.     

Spatial heterogeneity in soil particle size: does it affect the yield of plant communities with different species richness?

Aims Soil heterogeneity is ubiquitous in many ecosystems. We hypothesized that plant communities with higher species richness might be better adapted to soil heterogeneity and produce more biomass than those with lower richness. This is because there is niche differentiation among species and different species can complement each other and occupy a broader range of niches when plant species richness is high. However, no study has tested how soil particle heterogeneity affects the yield of plant communities, and whether such effects depend on the spatial scale of the heterogeneity and the species richness within the communities.Methods In a greenhouse experiment, we sowed seeds of four-species or eight-species mixtures in three heterogeneous treatments consisting of 32, 8 or 2 patches of both small (1.5mm) and large quartz (3.0mm) particles arranged in a chessboard manner and one homogeneous treatment with an even mixture of small and large quartz particles.Important findings Biomass production was significantly greater in the communities with high species richness than those with low species richness. However, soil particle heterogeneity or its interactions with patch scale or species richness did not significantly affect biomass production of the experimental communities. This work indicates that plant species richness may have a bigger impact on plant productivity than soil particle heterogeneity. Further studies should consider multiple sets of plant species during longer time periods to unravel the potential mechanisms of soil heterogeneity and its interactions with the impacts of species richness on community yield and species coexistence.     

Herbaceous plant richness and vegetation cover in Mediterranean grasslands and shrublands

Different types of relationship between herbaceous species richness and several parameters indicating abundance of plant material (herbaceous, woody plants, litter and bare ground cover) are presented. The data were obtained from 50 sites along a 300 km strip running from E to W within Spain and Portugal. Each site was representative of the silvo-pastoral landscape of the Mediterranean type ecosystems of the Iberian peninsula, and contained two neighboring patches, one of grassland and the other of shrubland. 3,600 20 × 20 cm subplots were randomly located (72 per site, 36 per patch) crossing the boundary grassland/shrubland. This approach allowed us to analyze the richness-occupation relationship of the space from different points of view: among and within the sites, and among and within the grassland and shrubland plant communities. We found a unimodal relationship between richness-cover similar to the one generally accepted between richness and biomass. Our results show that the dependence of this relationship varies depending on the spatial scale of the analysis and on the type of data used. When the whole region is taken into account, significant unimodal relationships are found between richness and herbaceous cover, litter and bare ground, and a negative linear relationship with woody plant cover. Within the sites there are mainly linear or non-significant relationships. But the results also depend on the type of communities analyzed. In pastures, the unimodal relationship represents the combination of positive and negative linear responses for low and high cover values, respectively. The value for herbaceous cover in which maximum richness occurs is around 60%. In shrublands, this value for cover also corresponds to maximum species richness, although the possibilities of reaching it are limited by other variables, such as woody plant cover. This implies that, on not considering variability at local scale, the relationship is linear and positive. This paper shows the existence of a common model related to herbaceous cover, but this model has multiple controlling factors that act differently in each type of community.