Determinants of avian species richness at different spatial scales

ABSTRACT. Studies of factors influencing avian biodiversity yield very different results depending on the spatial scale at which species richness is calculated. Ecological studies at small spatial scales (plot size 0.0025–0.4 km²) emphasize the importance of habitat diversity, whereas biogeographical studies at large spatial scales (quadrat size 400–50,000 km²) emphasize variables related to available energy such as temperature. In order to bridge the gap between those two approaches the bird atlas data set of Lake Constance was used to study factors determining avian species diversity at the intermediate spatial scales of landscapes (quadrat size 4–36 km²). At these spatial scales bird species richness was influenced by habitat diversity and not by variables related to available energy probably because, at the landscape scale, variation in available energy is small. Changing quadrat size between 4 and 36 km², but keeping the geographical extension of the study constant resulted in profound changes in the degree to which the amount of different habitat types was correlated with species richness. This suggests that high species diversity is achieved by different management regimes depending on the spatial scale at which species richness is calculated. However, generally, avian species diversity seems to be determined by spatial heterogeneity at the corresponding spatial scale. Thus, protecting the diversity of landscapes and ecosystems appears to ensure also high levels of species diversity.     

The Goldilocks effect: intermittent streams sustain more plant species than those with perennial or ephemeral flow

1. Drylands worldwide are typified by extreme variability in hydrologic processes, which structures riparian communities at various temporal and spatial scales. One key question is how underlying differences in hydrology over the length of interrupted perennial rivers influence spatial and temporal patterns in species richness and species composition. 2. We examined effects of differences in dry season hydrology on species richness, composition and cover of herbaceous plant communities in the streamside zone (the zone influenced directly by low flows in the channel). Data were collected at ephemeral, intermittent and perennial flow reaches on three rivers of the desert Southwest (Arizona, U.S.A.): Lower Cienega Creek, Hassayampa River and Lower San Pedro River. 3. Patterns of species richness varied with temporal scale of analysis, that is between single‐year and multi‐year time frames. At the annual timescale, quadrat species richness (m^?2) and herbaceous cover were higher at sites with perennial flow than at either intermittent or ephemeral sites. In contrast to this single‐year pattern, the highest long‐term richness occurred at intermittent sites. 4. Quadrat species richness, total species richness at a site (per 18 1‐m² plots) and cover were more variable year to year at non‐perennial sites than at perennial flow sites. On two of the three rivers, ephemeral sites had the highest inter‐annual compositional variance, while the perennial sites had the lowest. 5. Compositional differences between the hydrologic site types were dominated by species turnover, not nestedness. The perennial sites had more wetland and perennial species than the other two site types. The intermittent sites had more annual species than did the other two types. 6. High long‐term species richness and distinct species composition of intermittent sites are probably sustained by pronounced temporal variability in environmental conditions (i.e. frequent and persistent flow events, and dry periods). Plants at these sites take advantage of greater moisture than those at ephemeral sites and also experience less competition from resident species than those at perennial sites. 7. Conservation of desert riparian diversity depends upon the protection of consistently wet conditions at perennial flow sites, as well as the maintenance of the processes that cause fluctuations in environmental conditions at non‐perennial sites.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Broad‐scale patterns in plant diversity vary between land uses in a variegated temperate Australian agricultural landscape

Plant diversity is threatened in many agricultural landscapes. Our understanding of patterns of plant diversity in these landscapes is mainly based on small‐scale (<1000 m²) observations of species richness. However, such observations are insufficient for detecting the spatial heterogeneity of vegetation composition. In a case‐study farm on the North‐West Slopes of New South Wales, Australia, we observed species richness at four scales (quadrat, patch, land use and landscape) across five land uses (grazed and ungrazed woodlands, native pastures, roadsides and crops). We applied two landscape ecological models to assess the contribution of these land uses to landscape species richness: (i) additive partitioning of diversity at multiple spatial scales, and (ii) a measure of habitat specificity – the effective number of species that a patch contributes to landscape species richness. Native pastures had less variation between patches than grazed and ungrazed woodlands, and hence were less species‐rich at the landscape scale, despite having similar richness to woodlands at the quadrat and patch scale. Habitat specificity was significantly higher for ungrazed woodland patches than all other land uses. Our results showed that in this landscape, ungrazed woodland patches had a higher contribution than the grazed land uses to landscape species richness. These results have implications for the conservation management of this landscape, and highlighted the need for greater consensus on the influence of different land uses on landscape patterns of plant diversity.     

Multiscale Patterns of Riparian Plant Diversity and Implications for Restoration

Planning riparian restoration to resemble historic reference conditions requires an understanding of both local and regional patterns of plant species diversity. Thus, understanding species distributions at multiple spatial scales is essential to improve restoration planting success, to enhance long‐term ecosystem functioning, and to match restoration planting designs with historic biogeographic distributions. To inform restoration planning, we examined the biogeographic patterns of riparian plant diversity at local and regional scales within a major western U.S.A. drainage, California's Sacramento—San Joaquin Valley. We analyzed patterns of species richness and complementarity (β‐diversity) across two scales: the watershed scale and the floodplain scale. At the watershed scale, spatial patterns of native riparian richness were driven by herbaceous species, whereas woody species were largely cosmopolitan across the nearly 38,000 km² study area. At the floodplain scale, riparian floras reflected species richness and dissimilarity patterns related to hydrological and disturbance‐driven successional sequences. These findings reinforce the importance of concurrently evaluating both local and regional processes that promote species diversity and distribution of native riparian flora. Furthermore, as restoration activities become more prevalent across the landscape, strategies for restoration outcomes should emulate the patterns of species diversity and biogeographic distributions found at regional scales.     

不同植物类群物种丰富度垂直格局分形特征的比较

格局和过程一直是生态学的核心问题,该文应用“非布朗运动随机分形”模型,在北京东灵山远离人为干扰地区的阴坡设置了两条宽2 m的样带,描述和比较了暖温带落叶阔叶林区乔木、灌木、草本以及总的植物物种丰富度在不同尺度上沿海拔梯度变异特征,以及相关的生态过程。研究结果表明:1)直接梯度法刻画了植物物种丰富度在海拔梯度上的总体变化趋势,并不能从中获得更多的信息。“非布朗运动随机分形”模型揭示了不同尺度上物种丰富度的空间变化并能和生态过程相联系,研究表明该模型适合对乔木、灌木和草本植物物种丰富度的垂直梯度格局特征进行描述,但对总的植物物种丰富度有一定的局限性。2)乔木物种丰富度在海拔梯度上的变化主要受两种生态过程的控制,小尺度上分形维数接近于2,主要受独立的不具有长程相关的生态过程控制,呈近随机分布;大尺度上分形维数接近于1,主要受自相关范围大、具有长程关联特性的生态过程控制,随海拔上升呈近似单调下降的趋势。灌木和草本物种丰富度在海拔梯度上的变化主要受3种生态过程的控制,小尺度和大尺度上的特征与乔木相似;但在中等尺度上,分形维数接近于1.5,它们以近似布朗运动的形式变化。虽然乔木和灌木物种丰富度在海拔梯度上变化的尺度范围相似,但决定它们的主要生态过程完全不同,或相同的生态过程以截然不同的方式作用于两种不同的植物类群。总的植物物种丰富度在整个尺度范围内呈现标度不变性,这可能是由于决定其变化的生态过程作用尺度紧密相关,也可能因为该文研究范围的局限性。     

Grazing effects on plant functional group diversity in Mediterranean shrublands

Grazing is one of the prevalent human activities that even today are taking place inside protected areas with direct or indirect effects on ecosystems. In this study we analyzed the effects of grazing on plant species diversity, plant functional group (PFG) diversity and community composition of shrublands. We analyzed plant diversity data from 582 sampling plots located in 66 protected areas of the Greek Natura 2000 network, containing in total 1102 plant species and subspecies. We also classified a priori all plant species in seven PFGs: annual forbs, annual grasses/sedges, legumes, perennial forbs, perennial grasses/sedges, small shrubs and tall shrubs. For each site, grazing intensity was estimated in four classes (no grazing, low, medium and high grazing intensity). We found that, at the spatial and temporal scale of this study, as grazing intensity increased, so did total species richness. However, each PFG displayed a different response to grazing. Short-lived species (annual grasses or forbs and legumes) benefited from grazing and their species richness and proportion in the community increased with grazing. Perennial grasses and forbs species richness increased with grazing intensity, but their dominance decreased, since their proportion in the community declined. Short shrub species richness remained unaffected by grazing, while tall shrub diversity decreased. Finally, in sites without grazing the spatial pattern of species richness of the different PFGs was not congruent with each other, while in grazed sites they were significantly positively correlated (with the exception of tall shrubs). This finding may imply that grazing is a selective pressure organizing the community structure, and imposing a certain contribution of each PFG. So, in Mediterranean shrublands in protected areas with a long historical record of grazing, it seems that grazing promotes species diversity and its continuation on a portion of the landscape may be a necessary part of an effective management plan.     

Shrubs as ecosystem engineers in a coastal dune: influences on plant populations,communities and ecosystems

Question: How do two shrubs with contrasting life‐history characteristics influence abundance of dominant plant taxa, species richness and aboveground biomass of grasses and forbs, litter accumulation, nitrogen pools and mineralization rates? How are these shrubs – and thus their effects on populations, communities and ecosystems – distributed spatially across the landscape? Location: Coastal hind‐dune system, Bodega Head, northern California. Methods: In each of 4 years, we compared vegetation, leaf litter and soil nitrogen under canopies of two native shrubs –Ericameria ericoides and the nitrogen‐fixing Lupinus chamissonis– with those in adjacent open dunes. Results: At the population level, density and cover of the native forb Claytonia perfoliata and the exotic grass Bromus diandrus were higher under shrubs than in shrub‐free areas, whereas they were lower under shrubs for the exotic grass Vulpia bromoides. In contrast, cover of three native moss species was highest under Ericameria and equally low under Lupinus and shrub‐free areas. At community level, species richness and aboveground biomass of herbaceous dicots was lower beneath shrubs, whereas no pattern emerged for grasses. At ecosystem level, areas beneath shrubs accumulated more leaf litter and had larger pools of soil ammonium and nitrate. Rates of nitrate mineralization were higher under Lupinus, followed by Ericameria and then open dune. At landscape level, the two shrubs – and their distinctive vegetation and soils – frequently had uniform spatial distributions, and the distance separating neighbouring shrubs increased as their combined sizes increased. Conclusions: Collectively, these data suggest that both shrubs serve as ecosystem engineers in this coastal dune, having influences at multiple levels of biological organization. Our data also suggest that intraspecific competition influenced the spatial distributions of these shrubs and thus altered the distribution of their effects throughout the landscape.     

哀牢山亚热带中山湿性常绿阔叶林树种beta多样性格局形成的驱动力

Beta多样性通常指群落在时间和空间上物种组成的差异, 包括物种周转组分和物种丰富度差异组分。驱动beta多样性格局形成的生态过程决定了群落的时空动态, 然而关于beta多样性及其两个组分格局形成的驱动力还存在较多争议。以往研究表明, beta多样性的格局存在取样尺度的依赖性, 驱动其形成的生态过程在不同取样尺度下的相对重要性也随之改变。本研究以哀牢山亚热带中山湿性常绿阔叶林20 ha动态监测样地为研究对象, 在不同取样尺度上, 将样方间的Bray-Curtis指数分解为物种周转组分和物种丰富度差异组分, 通过典范冗余分析和方差分解的方法揭示环境过滤和扩散限制对于beta多样性及其两个组分格局形成的相对重要性及其尺度依赖性。结果表明: (1) beta多样性、物种周转组分和物种丰富度差异组分均随取样尺度的增大而减小。在不同取样尺度下, 物种周转组分对于beta多样性的贡献始终占主导地位。(2)随着取样尺度的增大, 环境过滤驱动beta多样性格局形成的相对重要性逐渐增加, 而扩散限制的相对重要性逐渐降低。本研究进一步证实了取样尺度在beta多样性格局形成及其驱动力定量评价中的重要性, 今后的研究需要进一步解析上述尺度效应的形成机制。     

Composition and diversity of herbaceous patches in woody vegetation: The effects of grazing,soil seed bank,patch spatial properties and scale

Herbaceous plants contribute much to plant diversity in Mediterranean-type ecosystems though mostly occupying relatively small patches within the dense woody vegetation. While studying species diversity in the herbaceous patches, we hypothesized that grazing, soil seed bank, and spatial properties of the patch affect plant diversity and composition at different spatial scales. The study site was in an LTER site located in the Mediterranean region in north Israel. We determined herbaceous species composition in: (1) randomly sampled quadrats in herbaceous patches in grazed and un-grazed plots; (2) soil seed bank samples taken from the same patches and germinated under optimal greenhouse conditions; (3) quadrats in the same patches sown with a homogenous mixture of local soil samples. Using GIS methods, we determined small-scale spatial characteristics of the herbaceous patches. Alpha and beta diversities were calculated at the patch and plot scales using Shannon's entropy H. Grazing increased alpha diversity of local untreated seed bank samples but decreased alpha diversity of the artificial homogenous soil seed bank mixture at both patch and plot scales. Positive relation between alpha diversity and patch area was detected only under grazing. Grazing increased beta diversity in all three treatments at the patch scale. Grazing decreased the similarity in species composition between above-ground vegetation and soil seed bank. The results indicate that moderate cattle-grazing affects species diversity in the herbaceous patches within the dense maquis. This effect is scale-dependent, and interacts with the effects of soil seed bank and patch spatial-properties: without grazing soil seed bank plays a more important role than patch spatial properties, but under grazing the size and the accessibility of the patch are more important in the determination of herbaceous species composition.     

Spatial variation of species diversity across scales in an old-growth temperate forest of China

Species richness and abundance are the two most important diversity variables. Species abundance is additive when aggregated across spatial scale, whereas species richness is non-additive. This study analyzes the effect of spatial scale and site on species abundance and richness in a 25-ha temperate forest plot in the Changbai Mountains, northeastern China. The result shows that species abundance and richness are not only dependent on spatial scales, but also dependent on site. Species abundance responds linearly to changes of spatial scale with no intersection in different sites of the study area. However, although species richness also increases with the increase of spatial scale, there are some intersections for the different sites, suggesting that a species-rich site does not always have a high value if the spatial scale is changed. In all, with respect to additive variables, it is relatively easy to extrapolate them from one spatial scale to another spatial scale, as they and the spatial scale usually form a linear relationship. In contrast, non-additive variables are difficult to extrapolate across spatial scales, because they often respond nonlinearly to spatial scale changes. In order to extrapolate these non-additive variables across spatial scales, it is necessary to estimate the relationships between them and spatial scales. As a result, extrapolation of information among spatial scales may be possible, but very difficult, especially for non-additive variables. Because the 25-ha Changbai plot is very small compared to the extent of the world temperate forests, and the vegetation is a relatively uniform type, more such studies in other ecosystems are needed before theories and generalization about scaling effects can be formulated.     

Cross-spatial-scale patterns in the facilitative effect of shrubs and potential for restoration of desert steppe

Facilitation (positive plant–plant interactions) is a potential means to accelerate vegetation restoration in arid areas. Shrubs can accelerate vegetation recovery by means of soil amelioration, but this effect has not been evaluated at large spatial scales or across scales. Here, we examined the facilitative function of shrub change across spatial scales at a desert steppe in Mongolia. Using a high-resolution satellite image, we established five 2500 m² plots in each of three shrub density classes (low, moderate, high) in a desert steppe in Mongolia. To evaluate the facilitative functions of shrubs at multiple spatial scales, we recorded the total number of plant species at three nested spatial scales in each plot: 25, 400, and 2500 m². The facilitative effect of shrubs on plant species richness was more pronounced at larger scales. Denser shrub communities increased plant species diversity at a larger scale. However, the increased taxonomic diversity was not clearly related to increased functional diversity in this system. This scale dependency in species diversity can be explained by the degree to which spatial heterogeneity of habitats within the plots increased as plot size increased. These results support the hypothesis of scale-dependent changes in the balance between facilitation and competition. Therefore, transplanting shrub saplings at high-density and a larger scale could potentially improve the success of vegetation restoration in arid regions.     

Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis

It has been suggested that a heterogeneous environment enhances species richness and allows for the coexistence of species. However, there is increasing evidence that environmental heterogeneity can have no effect or even a negative effect on plant species richness and plant coexistence at a local scale. We examined whether plant species richness increases with local heterogeneity in the water table depth, microtopography, pH and light availability in a swamp forest community at three local spatial scales (grain: 0.6, 1.2 and 11.4 m). We also used the variance partitioning approach to assess the relative contributions of niche-based and other spatial processes to species occurrence. We found that heterogeneity in microtopography and light availability positively correlated with species richness, in accordance with the habitat heterogeneity hypothesis. However, we recorded different heterogeneity-diversity relationships for particular functional species groups. An increase in the richness of bryophytes and woody plant species was generally related to habitat heterogeneity at all measured spatial scales, whereas a low impact on herbaceous species richness was recorded only at the 11.4 m scale. The distribution of herbaceous plants was primarily explained by other spatial processes, such as dispersal, in contrast to the occurrence of bryophytes, which was better explained by environmental factors. Our results suggest that both niche-based and other spatial processes are important determinants of the plant composition and species turnover at local spatial scales in swamp forests.     

The effect of overstorey proteas on plant species richness in South African mountain fynbos

Two South African mountain fynbos sites, similar in drainage, elevation, slope angle, slope aspect and soil type but with differing fire histories, were studied to measure how the effect of high densities of overstorey proteas in one fire cycle affects the α-diversity levels of the plant community in the following fire-cycle, how their repeated absence due to several short fire-cycles affects their species richness and finally, at what spatial scale such patterns are most appropriately measured. High prefire canopy cover percentages and densities of overstorey proteas increase the postfire α-diversity of understorey species. In addition, the increase in species richness observed occurred for all higher plant life history types present. At sites where one or more short fire cycles resulted in the repeated absence of overstorey proteas, the number of plant species present in the understorey was lower than at a site where overstorey proteas persisted. These results are dependent on the spatial scale at which the α-diversity of understorey species is measured. At small quadrat sizes (< 5 m²), overstorey proteas decrease the number of understorey species present, while at larger quadrat sizes (100 m²) higher species richness is observed. The contradiction in conclusions when α-diversity is measured at different spatial scales can be attributed to the patchiness of fynbos communities. Overstorey proteas play an important role in maintaining the patchiness component of fynbos communities by diminishing the effect of understorey resprouting species, making available regeneration niches for the maintenance of plant species richness. Where small quadrats are used, the effect of patchiness on the dynamics of the mountain fynbos community is lost. Thus, it is the fire history prior to the last fire and how it affects overstorey proteas that is important in the determination of α-diversity levels in mountain fynbos plant communities.     

Ecological stoichiometry explains larger-scale facilitation processes by shrubs on species coexistence among understory plants

Plant facilitation (positive plant–plant interactions) strongly influences biodiversity, structure, and dynamics in plant communities, and the topic has received considerable attention among ecologists. Most studies of facilitation processes by shrubs have been conducted at small spatial scales between shrubs and their neighboring species. Yet, we know little about whether facilitation processes by shrubs at a small scale (i.e., a patch scale) also work at a larger scale (i.e., a site scale) in terms of the maintenance of biodiversity. Here, we report that the facilitative effects of shrubs on plant diversity at a larger scale can be explained by changing ecological stoichiometry. The soil fertility showed unimodal shape along shrub cover gradient, suggesting that the facilitative effects of a shrub do not necessarily increase as the shrub develops. The unimodal shape of dependence of plant species richness on shrub cover probably was generated by the unimodal dependence of soil fertility on shrub cover. Soil nutrient enrichment by shrubs shifted low N:P ratios of plant communities with low levels of shrub cover to more balanced N:P ratios at intermediate levels of shrub cover. At the peak N:P ratio along the gradient in shrub cover, the maximum species richness and functional richness were observed, which was consistent with the unimodal relationship predicted by the resource balance hypothesis. Thus, our findings showed that facilitation processes by shrubs at a patch scale also work at a larger scale in terms of the maintenance of biodiversity. Because observed larger-scale facilitation processes are enhanced at some intermediate levels of shrub cover, this study offers practical insight into the need for management practices that allow some intermediate levels of grazing by livestock for optimizing the role of larger-scale facilitation processes in the maintenance of biodiversity and ecosystem functioning in arid and semi-arid rangelands.     

Regional and local influence of grazing activity on the diversity of a semi-arid dung beetle community

This study analyses the effect of resource availability (i.e. sheep dung) on dung beetle communities in an arid region of Central Spain, both at regional and at local scales. A total of 18 sites within 600 km² were sampled for the regional analysis and 16 sites within the 30 km² of an Iberian municipality were sampled for the local analysis. Spatial and environmental characteristics of sampling sites were also compiled at both scales, including measures of grazing activity (livestock density at regional scale, and two counts of rabbit and sheep dung at local scale). At a regional scale, any environmental or spatial variable can help to explain the variation in abundance. However, species richness was related to summer precipitation and composition was related to elevation. At local scale, abundance is not significantly related to any of the environmental variables, but species richness was related to the local amount of sheep dung (27% of variance). The amount of dung in a 2‐km buffer around the site accounts for 27–32% of variance in abundance and 60–65% of variance in species richness. The presence of the flock with the highest sheep density explains 53% of abundance variability and 73% of species richness variance. A cluster analysis of localities identified two main groups, one characterized by a lower abundance and species richness that can be considered a nested subsample of the species‐rich group. The mean and maximum amount of sheep dung in the sites separated by less than 2 km are the only significant explanatory variables able to discriminate both groups. These results suggest that grazing intensity (and the associated increase in the amount of trophic resources) is a key factor in determining local variation in the diversity and composition of dung beetle assemblages. However, dung beetle assemblages are not spatially independent at the analysed resolution, and the amount of dung in the surroundings seems to be more important for locally collected species than the dung effectively found in the site. Although differences in the availability and quantity of trophic resources among nearby sites could be affecting the population dynamics and dispersion of dung beetles within a locality, sites with larger populations, and greater species numbers would not be able to exercise enough influence as to bring about a complete local faunistic homogenization.     

Biogeography of vascular plants on habitat islands, peninsulas and mainlands in an eascentral Swedish agricultural landscape

The increase of island species richness with area can be explained by an increase in habitat diversity or by an equilibrium of species immigration and extinction. We examined vascular plant species richness in 39 sites (24 habitat islands, 7 'habitat peninsulas' and 8 comparable 'mainland' sites). We sampled at three scales: whole sites, meadows within sites and quadrats (4 m × 4 m) within meadows. All sites (10–10⁴ m²) contained natural vegetation within arable fields in eascentral Sweden. There was a strong correlation between species richness and area for whole sites and for meadows There was no correlation, however, between species richness in quadrats and site area. The difference between site and meadow results on one side and quadrat results on the other suggests that species richness increases with whole site area primarily because large sites are more diverse than smaller ones. Speciearea relationships did not differ between islands, peninsula and mainland sites. Thus, patterns of species richness on our sites were more consistent with habitat diversity than an immigratioextinction equilibrium.     

Distribution of Vascular Plant Species Richness Along an Elevational Gradient in the Dongling Mountains, Beijing, China

Quantifying spatial patterns of species richness and determining the processes that give rise to these patterns are core problems In blodlveralty theory. The aim of the present paper was to more accurately detect patterns of vascular species richness at different scales along altitudinal gradients in order to further our understanding of biodlverslty patterns and to facilitate studies on relationships between biodiversity and environmental factors. Species richness patterns of total vascular plants species, including trees, shrubs, and herbs, were measured along an altitudinal gradient on one transect on a shady slope in the Dongling Mountains, near Beijing,China. Direct gradient analysis, regression analysis, and geostatistics were applied to describe the spatial patterns of species richness. We found that total vascular species richness did not exhibit a linear pattern of change with altitude, although species groups with different ecological features showed strong elevational patterns different from total species richness. In addition to total vascular plants, analysis of trees, shrubs, and herbs demonstrated remarkable hierarchical structures of species richness with altitude (i.e. patchy structures at small scales and gradients at large scales). Species richness for trees and shrubs had similar spatial characteristics at different scales, but differed from herbs. These results indicated that species groups with similar ecological features exhibit similar biodlveraity patterns with altitude, and studies of biodiversity based on species groups with similar ecological properties or life forms would advance our understanding of variations in species diversity. Furthermore, the gradients or trends appeared to be due mainly to local variations in species richness means with altitude. We also found that the range of spatial scale dependencies of species richness for total vascular plants, trees, shrubs, and herbs was relatively large. Thus, to detect the relationships betweenspecies richness with environmental factors along altitudinal gradients, it was necessary to quantify the scale dependencies of environmental factors in the sampling design or when establishing non-linear models.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.