Benthic microalgal diversity enhanced by spatial heterogeneity of grazing

This study presents model experiments on the effect of the spatial pattern of herbivory on primary producer diversity. Microalgal biofilms (periphyton) were exposed to different mixtures of two benthic herbivores, the isopod Idothea chelipes and the gastropod Littorina littorea. The herbivores are similar in their feeding selectivity but differ strongly in the spatial pattern of grazing. Idothea did not increase the spatial heterogeneity of algal cell densities beyond the level of ungrazed controls (<1 order of magnitude between local minima and maxima at the 1 mm² scale). Littorina grazing, in contrast, created a pronounced spatial heterogeneity with maximum:minmum ratios of almost 3 orders of magnitude. When algae were exposed to mixtures of both gazers, the spatial heterogeneity of microalgal cell densities increased with an increasing proportion of Littorina in the herbivore mixture. Algal species richness, diversity and evenness also increased with increasing proportions of Littorina, and was highly significantly correlated with the spatial heterogeneity of cell densities. Received: 25 May 1999 / Accepted: 14 September 1999     

Spatial patterns of species distributions in grazed subalpine grasslands

Spatial patterns of species diversity have important influences on the functioning of ecosystems, and the effect of livestock grazing on spatial heterogeneity can differ depending on the scale of the analysis. This study examined the effects of grazing on the spatial patterns of species distributions and whether the effects of grazing on the spatial distributions of a species vary with the scale of the analysis. Data were collected at three locations in the subalpine grasslands of Ordesa-Monte Perdido National Park and Aísa Valley, Central Pyrenees, Spain, which differed in mean stocking rates. Aspect explained about one-third of the environmental variation in species distributions. In flat areas, spatial variation in species composition varied with grazing intensity at two scales. At a coarse scale (among vegetation patches), grazing promoted patchiness, and among-transect variation in species diversity and grazing intensity were positively correlated. At a fine scale (within vegetation patches), the disruption of the self-organizing processes of the species spatial distributions resulted in a reduction in the long-range spatial autocorrelations of some of the characteristic species and in the homogenization of species spatial distributions. The presence of encroaching Echinospartum horridum had a significant influence on the effect of grazing on south-facing grassland slopes.     

Effects of large herbivores on grassland arthropod diversity

Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio‐temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top‐down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.     

放牧对草地昆虫多样性的影响研究进展

在草地生态系统中,大型草食动物放牧是重要的管理方式之一,对草地生物多样性起着关键的驱动作用。昆虫是草地生态系统中生物多样性的重要组成成分,对生态系统的食物网结构以及其功能与稳定性起着关键作用。已有研究结果表明,大型草食动物与昆虫存在密切联系,放牧对草地昆虫多样性或有正向、或负向、或无明显作用,这依赖于放牧管理方式、昆虫类群以及草地类型。放牧必然通过直接(采食、践踏或粪尿)或间接(植物群落组成或植被结构)作用对昆虫多样性产生显著的影响。当前,关于大型草食动物放牧对草地昆虫多样性影响研究较多,但是,从研究系统性、深入性和延续性来说还存在一定问题。本文在综述国内外对放牧对草地昆虫多样性的影响研究基础上,提出了今后的研究方向,对于理解放牧管理的草地昆虫多样性变化规律,以及为积极探索维持草地昆虫多样性的长期有效的科学管理措施提供理论指导。     

Grazing and productivity alter individual grass size dynamics in semi-arid woodlands

The spatial arrangement of perennial vegetation is critical for ecosystem function in drylands. While much is known about how vegetation patches respond to grazing and abiotic conditions, the size dynamics of individual plants is mostly limited to theoretical studies. We measured the size distribution (mean, variance, skewness) and density of individual grasses, and grass species composition at 451 sites spanning a range of grazing intensities across three broad vegetation communities in semi-arid eastern Australia. We assessed the relative role of grazing by livestock (cattle and sheep), native (kangaroos) and introduced (rabbits) free ranging herbivores, and several environmental measures (productivity, diversity, composition and groundstorey plant cover) on the size distribution and density of individual grasses. We found mean grass size and density were more sensitive to shifts in grazing intensity and environmental conditions than size variance or the frequency of the smallest individuals (skewness), and shifts were mostly driven by site productivity and cattle and kangaroo grazing. Sheep grazing only reduced mean grass size, and rabbit grazing had no consistent effects. Importantly, we found that site productivity and species composition altered the impacts of grazing on grass density and size distribution. For example, increasing cattle grazing led to larger grasses in low productivity sites. It also led to larger, denser, more variable-sized grasses among grass species from sites with finer soil texture. Increasing kangaroo grazing led to smaller, denser individuals among grass species from sites with coarse soil texture. At high diversity sites kangaroo grazing led to denser, more homogenised grass sizes with a lower frequency of small individuals. Understanding the in situ response of individual plant sizes gives us insights into the processes driving shifts in perennial vegetation patchiness, improving our ability to predict how the spatial arrangement of ecosystems might change under global change scenarios.     

Is livestock grazing compatible with amphibian diversity in the High Mountains of Córdoba, Argentina?

The High Mountains of Córdoba, Argentina have a long evolutionary history of grazing by large herbivores. However, about 400?years ago, European livestock were introduced and gradually replaced native herbivores. Since the 1920s, domestic herbivores have been the only large herbivores present in the area, causing severe soil erosion and a threat to the system diversity. The endemic fauna of the region includes four amphibian species. We evaluated the effect of livestock rearing on amphibian diversity and water bodies in woodlands and grasslands of the High Mountains of Córdoba. The work was conducted on stream stretches and ponds in two contrasting grazing situations: an area with livestock presence and another area where livestock was excluded 14?years ago. The following variables were recorded at each sampling site: amphibian richness and abundance, percentage of emergent, submerged and peripheral vegetation in areas surrounding the water bodies, water pH, and water dissolved O₂. No significant differences were detected in amphibian diversity between streams of both grazing situations, whereas a greater diversity (p?<?0.01) was observed in ponds in grazed grasslands. Our results suggest that livestock rearing, qualitatively measured as grazing and 14?years of livestock exclusion, in the study area would not have negative effects on amphibian diversity. This finding might be due to the long evolutionary grazing history of the area, large-scale livestock exclusion exhibiting a novel scenario. The ongoing reintroduction of native grazers may provide the benefits of grazing without the consequent soil erosion and habitat degradation associated with domestic livestock.     

Patchiness in a minimal nutrient — phytoplankton model

We present a minimal two-component model that can exhibit various types of spatial patterns including patchiness. The model, comprising nutrients and phytoplankton, includes the effect of nutrient uptake by phytoplankton as a Holling type II functional response, and also includes the effect of zooplankton grazing on phytoplankton as a Holling type II non-dynamical term. The mean-field model without the diffusion and advection terms shows both bistability and limit-cycle oscillations as a few parameters such as the input rate of nutrients and the maximum feeding rate of zooplankton are changed. If the parameter values are chosen from the limit-cycle oscillation region, the corresponding reaction-advection-diffusion equations show spatial pattern formations by the combined effects of advection and diffusion by turbulent stirring and mixing, and biological interactions. As the nutrient input is increased, the system behaviour changes from the extinction of the entire phytoplankton to the formation of filamentous patterns, patchiness patterns and homogeneous distributions. These observations suggest that the spatial pattern of phytoplankton can function as an indicator to evaluate the eutrophication level in aquatic ecosystems.     

Herbivory and drought interact to enhance spatial patterning and diversity in a savanna understory

The combination of abiotic stress and consumer stress can have complex impacts on plant community structure. Effective conservation and management of semi-arid ecosystems requires an understanding of how different stresses interact to structure plant communities. We explored the separate and combined impacts of episodic drought, livestock grazing, and wild ungulate herbivory on species co-occurrence and diversity patterns in a relatively productive, semi-arid Acacia savanna. Specifically, we analyzed 9 years of biannual plant community data from the Kenya long-term exclosure experiment, a broad-scale manipulative experiment that has excluded different combinations of large mammalian herbivores from 18 4-ha plots since 1995. During droughts, we observed low species diversity and random species co-occurrence patterns. However, when rain followed a major drought, areas exposed to moderate cattle grazing displayed high species diversity and evidence of significant species aggregation. These patterns were not apparent in the absence of cattle, even if other large herbivores were present. To explore possible mechanisms, we examined patterns separately for common and rare species. We found that aggregation patterns were likely driven by rare species responding similarly to the availability of open micro-sites. Our results indicate that in a productive, fire-suppressed savanna, the combination of periodic drought and moderate cattle grazing can enhance plant biodiversity and fine-scale spatial heterogeneity by opening up space for species that are otherwise rare or cryptic. Our findings also emphasize that domestic herbivores can have significantly stronger impacts on plant community dynamics than wild herbivores, even in an ecosystem with a long history of grazing.     

Spatial heterogeneity and plant species richness at different spatial scales under rabbit grazing

Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.     

Effects ofFucus vesiculosus covering intertidal mussel beds in the Wadden Sea

The brown algaFucus vesiculosus formamytili (Nienburg) Nienhuis covered about 70% of mussel bed (Mytilus edulis) surface area in the lower intertidal zone of Königshafen, a sheltered sandy bay near the island of Sylt in the North Sea. Mean biomass in dense patches was 584 g ash-free dry weight m^?2 in summer. On experimental mussel beds, fucoid cover enhanced mud accumulation and decreased mussel density. The position of mussels underneath algal canopy was mainly endobenthic (87% of mussels with >1/3 of shell sunk into mud). In the absence of fucoids, mussels generated epibenthic garlands (81% of mussels with <1/3 of shell buried in mud). Mussel density underneath fucoid cover was 40 to 73% of mussel density without algae. On natural beds, barnacles (Balanidae), periwinkles (Littorina littorea) and crabs (particularly juveniles ofCarcinus maenas) were significantly less abundant in the presence of fucoids, presumably because most of the mussels were covered with sediment, whereas in the absence of fucoids, epibenthic mussel clumps provided substratum as well as interstitial hiding places. The endobenthic macrofauna showed little difference between covered and uncovered mussel beds. On the other hand, grazing herbivores — the flat periwinkleLittorina mariae, the isopodJaera albifrons and the amphipodsGammarus spp. — were more abundant at equivalent sites with fucoid cover. The patchy growth ofFucus vesiculosus on mussel beds in the intertidal Wadden Sea affects mussels and their epibionts negatively, but supports various herbivores and increases overall benthic diversity.     

An approach based on the total‐species accumulation curve and higher taxon richness to estimate realistic upper limits in regional species richness

Most of accumulation curves tend to underestimate species richness, as they do not consider spatial heterogeneity in species distribution, or are structured to provide lower bound estimates and limited extrapolations. The total‐species (T–S) curve allows extrapolations over large areas while taking into account spatial heterogeneity, making this estimator more prone to attempt upper bound estimates of regional species richness. However, the T–S curve may overestimate species richness due to (1) the mismatch among the spatial units used in the accumulation model and the actual units of variation in β‐diversity across the region, (2) small‐scale patchiness, and/or (3) patterns of rarity of species. We propose a new framework allowing the T–S curve to limit overestimation and give an application to a large dataset of marine mollusks spanning over 11 km² of subtidal bottom (W Mediterranean). As accumulation patterns are closely related across the taxonomic hierarchy up to family level, improvements of the T–S curve leading to more realistic estimates of family richness, that is, not exceeding the maximum number of known families potentially present in the area, can be considered as conducive to more realistic estimates of species richness. Results on real data showed that improvements of the T–S curve to accounts for true variations in β‐diversity within the sampled areas, small‐scale patchiness, and rarity of families led to the most plausible richness when all aspects were considered in the model. Data on simulated communities indicated that in the presence of high heterogeneity, and when the proportion of rare species was not excessive (>2/3), the procedure led to almost unbiased estimates. Our findings highlighted the central role of variations in β‐diversity within the region when attempting to estimate species richness, providing a general framework exploiting the properties of the T–S curve and known family richness to estimate plausible upper bounds in γ‐diversity.     

Spatial heterogeneity as a multiscale characteristic of zooplankton community

Zooplankton spatial heterogeneity has profound effects on understanding and modelling of zooplankton population dynamics and interactions with other planktonic compartments, and consequently, on the structure and function of planktonic ecosystems. On the one hand, zooplankton heterogeneity at spatial and temporal scales of ecological interest is an important focus of aquatic ecology research because of its implications in models of productivity, herbivory, nutrient cycling and trophic interactions in planktonic ecosystems. On the other hand, estimating zooplankton spatial variation at the scale of an ecosystem, is a powerful tool to achieve accurate sampling design. This review concentrates on the spatial heterogeneity of marine and freshwater zooplankton with respect to scale. First to be examined are the concept of spatial heterogeneity, the sampling and statistical methods used to estimate zooplankton heterogeneity, and the scales at which marine and freshwater zooplankton heterogeneity occurs. Then, the most important abiotic and biotic processes driving zooplankton heterogeneity over a range of spatial scales are presented and illustrated by studies conducted over large and fine scales in both oceans and lakes. Coupling between abiotic and biotic processes is finally discussed in the context of the multiple driving forces hypothesis.Studies of zooplankton spatial heterogeneity refer both to the quantification of the degree of heterogeneity (measured heterogeneity) and to the estimation of the heterogeneity resulting from the interactions between the organisms and their environment (functional heterogeneity) (Kolasa & Rollo, 1991). To resolve the problem of measuring zooplankton patchiness on a wide range of spatial scales, advanced technologies (acoustic devices, the Optical Plankton Counter (OPC), and video systems) have been developed and tested in marine and freshwater ecosystems. A comparison of their potential applications and limitations is presented. Furthermore, many statistical tools have been developed to estimate the degree of measured heterogeneity; the three types most commonly used are indices of spatial aggregation, variance: mean ratio, and spatial analysis methods. The variance partitioning method proposed by Borcard et al. (1992) is presented as a promising tool to assess zooplankton functional heterogeneity.Nested patchiness is a common feature of zooplankton communities and spatial heterogeneity occurs on a hierarchical continuum of scales in both marine and freshwater environments. Zooplankton patchiness is the product of many physical processes interacting with many biological processes. In marine systems, patterns of zooplankton patchiness at mega- to macro-scales are mostly linked to large advective vectorial processes whereas at coarse-, fine- and micro-scales, physical turbulence and migratory, reproductive and swarm behaviors act together to structure zooplankton distribution patterns. In freshwater environments, physical advective forces related to currents of various energy levels, and vertical stratification of lake interact with biological processes, especially with vertical migration, to structure zooplankton community over large to fine- and micro-scales. Henceforth, the zooplankton community must be perceived as a spatially well-structured and dynamic system that requires a combination of both abiotic and biotic explanatory factors for a better comprehension and more realistic and reliable predictions of its ecology.     

Interactions between spatially separated herbivores indirectly alter plant diversity

Above‐ and belowground herbivores promote plant diversity when selectively feeding on dominant plant species, but little is known about their combined effects. Using a model system, we show that neutral effects of an aboveground herbivore and positive effects of a belowground herbivore on plant diversity became profoundly negative when adding these herbivores in combination. The non‐additive effects were explained by differences in plant preference between the aboveground‐ and the belowground herbivores and their consequences for indirect interactions among plant species. Simultaneous exposure to aboveground‐ and belowground herbivores led to plant communities being dominated by a few highly abundant species. As above‐ and belowground invertebrate herbivores generally differ in their mobility and local distribution patterns, our results strongly suggest that aboveground–belowground interactions contribute to local spatial heterogeneity of diversity patterns within plant communities.     

Physical Heterogeneity Increases Biofilm Resource Use and Its Molecular Diversity in Stream Mesocosms

Background

Evidence increasingly shows that stream ecosystems greatly contribute to global carbon fluxes. This involves a tight coupling between biofilms, the dominant form of microbial life in streams, and dissolved organic carbon (DOC), a very significant pool of organic carbon on Earth. Yet, the interactions between microbial biodiversity and the molecular diversity of resource use are poorly understood.

Methodology/Principal Findings

Using six 40-m-long streamside flumes, we created a gradient of streambed landscapes with increasing spatial flow heterogeneity to assess how physical heterogeneity, inherent to streams, affects biofilm diversity and DOC use. We determined bacterial biodiversity in all six landscapes using 16S-rRNA fingerprinting and measured carbon uptake from glucose and DOC experimentally injected to all six flumes. The diversity of DOC molecules removed from the water was determined from ultrahigh-resolution Fourier Transform Ion Cyclotron Resonance mass spectrometry (FTICR-MS). Bacterial beta diversity, glucose and DOC uptake, and the molecular diversity of DOC use all increased with increasing flow heterogeneity. Causal modeling and path analyses of the experimental data revealed that the uptake of glucose was largely driven by physical processes related to flow heterogeneity, whereas biodiversity effects, such as complementarity, most likely contributed to the enhanced uptake of putatively recalcitrant DOC compounds in the streambeds with higher flow heterogeneity.

Conclusions/Significance

Our results suggest biophysical mechanisms, including hydrodynamics and microbial complementarity effects, through which physical heterogeneity induces changes of resource use and carbon fluxes in streams. These findings highlight the importance of fine-scale streambed heterogeneity for microbial biodiversity and ecosystem functioning in streams, where homogenization and loss of habitats increasingly reduce biodiversity.     

不同放牧强度下短花针茅荒漠草原植被的空间异质性

分析不同放牧强度下植物群落中物种的空间分布特征, 有助于阐明群落在放牧胁迫下的演替规律。该研究基于幂函数法则, 探讨了不同放牧强度下短花针茅(Stipa breviflora)荒漠草原群落植物的频率和空间异质性。结果表明: 不同放牧强度下物种空间分布与幂函数法则能很好地吻合; 不同物种空间异质性具有特异性, 随着放牧强度的增加, 提高群落空间异质性的物种分别由无芒隐子草(Cleistogenes songorica)、冷蒿(Artemisia frigida)、短花针茅、银灰旋花(Convolvulus ammannii)等多个物种逐渐转变为以无芒隐子草、短花针茅为主的少数物种, 同时, 物种空间异质性大于群落空间异质性的物种数逐渐减少。     

科尔沁沙质草地植物群落高度空间异质性对不同放牧方式的响应

放牧是影响草地植被空间异质性的重要生物驱动因素。大型草食动物会通过采食、践踏和排泄粪便等方式影响草地植被空间异质性。同时,植物群落高度是重要的草地结构参数,但目前关于不同放牧方式对草地植物群落高度空间异质性的研究较少。该研究基于传统及地统计学方法,设置未放牧(NG)、牛单牧(CG)、羊单牧(SG)、牛羊混牧(MG) 4种处理,探究科尔沁沙质草地植物群落高度空间异质性对不同放牧方式的响应。结果表明:研究样地内植物群落平均高度为SG (38.86 cm)> NG(21.01 cm)> MG (17.29 cm)> CG (13.36 cm)。各处理下变异系数的结果均为中等变异,且变异范围在37.82%–66.97%之间,依次为NG> MG> CG> SG。通过对半方差函数参数进行最适模型拟合, NG、MG、CG、SG 4种处理分别对应球状模型、指数模型、线性模型、线性模型,且结构比大小为MG (82.7%)> NG (80.3%)> SG (40.2%)> CG (39.1%)。此外,采用Kriging插值法和分形维数对研究样地内植物群...     

Testing the role of biotic stress in the stress gradient hypothesis. Processes and patterns in arid rangelands

Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.     

Red deer mediate spatial and temporal plant heterogeneity in boreal forests

Selective herbivory can influence both spatial and temporal vegetation heterogeneity. For example, many northern European populations of free-ranging ungulates have reached unprecedented levels, which can influence plant species turnover, long-term maintenance of biodiversity and the subsequent stability of boreal ecosystems. However, the mechanisms by which large herbivores affect spatial and temporal vegetation heterogeneity remain poorly understood. Here, we combined a 10-year exclusion experiment with a herbivore intensity gradient to investigate how red deer (Cervus elaphus) acts as a driver of temporal and spatial heterogeneity in the understory of a boreal forest. We measured the two dimensions of heterogeneity as temporal and spatial species turnover. We found that temporal heterogeneity was positively related to herbivory intensity, and we found a similar trend for spatial heterogeneity. Removing red deer (exclosure) from our study system caused a distinct shift in species composition, both spatially (slow response) and temporally (quick response). Vegetation from which red deer had been excluded for 10 years showed the highest spatial heterogeneity, suggesting that the most stable forest understory will occur where there are no large herbivores. However, excluding red deer resulted in lower species diversity and greater dominance by a low number of plant species. If both stable but species rich ecosystems are the management goal, these findings suggest that naturally fluctuating, but moderate red deer densities should be sustained.     

The persistent spatial patchiness of limpet grazing

The characteristic variability of grazing has potential consequences for intertidal productivity and community structure, particularly as many of the underlying functional relationships are thought to be non linear. As a first approximation, it can be hypothesised that grazing is patchy over short time periods before a more uniform coverage is established over longer time scales. This prediction is supported by relatively short term observations previously made of limpet foraging. We used eight arrays of wax disks on each of four shores to test the hypothesis that grazing is patchy in the short term, but that this pattern is lost as the pattern of grazing averages out over longer time scales. Wax disk arrays were exposed for two weeks at a time for a period of six weeks in 2001 and in 2002 using the same set of disk holes each time. Grazing at the same disk location could therefore be measured over two weeks and over longer periods by averaging successive deployments. We used all three successive deployments to estimate the average grazing at each disk location over a six week period in 2001 and 2002. All six deployments were used to characterise the pattern of grazing at longer time scales. The spatial pattern of grazing in arrays was summarized using semivariogram analyses. For two-week deployments, the average standardized semivariance of grazing for disks separated by 20 cm was less than one. This pattern implies spatial autocorrelation of grazing at this scale. There was no support for the hypothesis that small scale patchiness in grazing would disappear over time. The average strength of spatial autocorrelation increased when data were integrated over longer periods. A preliminary analysis indicated that the degree of autocorrelation within arrays increased with grazing intensity at short time scales. Surface roughness disrupted autocorrelation of grazing over both short and long time scales. The persistent patchiness of grazing is likely to have implications for biofilm productivity, particularly on smoother shores.     

放牧和刈割条件下草山草坡群落空间异质性分析

采用变异矩分析和分形方法,研究了草山草坡群落在放牧和刈割条件下的空间异质性及空间自相关性,结果表明,群落空间格局有尺度依赖性,刈割条件下空物异质性及空间相关性弱,多样性梯度即β多样性小,放牧消除地形引起的样地差异,因而使空间异质性简单化。