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.     

Holocene dynamics of the alpine timberline in the High Sudetes

The study focuses on the High Sudetes that represent the most distinctive islands of alpine forest-free area among hercynian mid-mountains of Central Europe. Based on data from newly taken cores and previously published pollen profiles, comparison of the development of the alpine timberline position is carried out. The first of the analysed pollen profiles — the Labsky důl core in the Krkonoše Mts spans the whole period of the Holocene, the Keprník and Mezikotlí profiles in the Hruby Jeseník Mts bring information from the Subboreal/older Subatlanticum turn to the present. An exceptional position of the Krkonoše Mts in terms of permanent presence of the alpine belt throughout the Holocene was confirmed. Three oscillations of the alpine timberline during the Lower Holocene were detected in the profile from the Labsky důl site. In the Hruby Jeseník Mts a temperature dependent forest-free area existed at least since Subboreal to the present.     

The influence of scale and species pool on the relationship between vascular plant species richness and cover in an alpine area in Norway

Recent studies emphasise the potential importance of scale and species pool on the humped-back or unimodal relationship between species richness and productivity. We use a classic phytosociological data-set from Rondane, central south Norway, to evaluate the relative importance of these factors in an alpine area. The effect of species pool is assessed using plot scores from a Correspondence Analysis (CA) of the data. Generalised Additive Models (GAM) are used to relate vascular plant species richness to cover of vascular plants, CA plot scores, and plot area in different combinations. Species richness of vascular plants is unimodally related to total vascular plant cover. Plot scores of the first three CA axes (representing the effect of species pool) have a complex relationship with species richness, but explain a large fraction of the total deviance in richness. A humped relationship between richness and cover remains after accounting for CA plot scores in the model, i.e. the relationship is independent of species pool. The results suggest that the relationship between richness and cover changes from one vegetation type to another, as evaluated statistically through the importance of the interaction between cover and CA scores in explaining variation in richness. Plot area also influences the relationship. A unimodal relationship is only evident when small plot sizes are used, whereas a monotonically increasing relationship is found at large plot sizes. Plot area has the strongest effect on the unimodal relationship between richness and cover, whereas vegetation type has only a minor effect on this relationship. This revised version was published online in June 2006 with corrections to the Cover Date.     

The effect of area on local and regional elevational patterns of species richness

Aim To calculate the degree to which differences between local and regional elevational species richness patterns can be accounted for by the effects of regional area. Location Five elevational transects in Costa Rica, Ecuador, La Réunion, Mexico and Tanzania. Methods We sampled ferns in standardized field plots and collated regional species lists based on herbarium and literature data. We then used the Arrhenius function S = cA^z to correct regional species richness (S) for the effect of area (A) using three slightly different approaches, and compared the concordance of local and regional patterns prior to and after accounting for the effect of area on regional richness using linear regression analyses. Results We found a better concordance between local and regional elevational species richness after including the effect of area in the majority of cases. In several cases, local and regional patterns are very similar after accounting for area. In most of the cases, the maximum regional richness shifted to a higher elevation after accounting for area. Different approaches to correct for area resulted in qualitatively similar results. Main conclusions The differences between local and regional elevational richness patterns can at least partly be accounted for by area effects, suggesting that the underlying causes of elevational richness patterns might be the same at both spatial scales. Values used to account for the effect of area differ among the different study locations, showing that there is no generally applicable elevational species–area relationship.     

Macro‐scale bird species richness patterns of the East Asian mainland and islands: energy,area and isolation

Aim To create a map of bird species richness (BSR) in East Asia and to examine the effect of area, isolation, primary productivity, topographic heterogeneity, and human population density on BSR. Location East Asia (from 70° E to 180° E longitude), including the eastern half of the Palaearctic Region, the entire Oriental Region, and the entire Wallacea Subregion. Methods The breeding ranges of 2406 terrestrial bird species were mapped and overlaid to create a species richness map. The BSR map was transformed into a 100 × 100 km quadrat system, and BSR was analysed in relation to land area, average normalized difference vegetation index (NDVI), elevation range, and average population density. Results In general, BSR declined from the Tropics to the Arctic. In mainland East Asia, however, BSR was highest around the Tropic of Cancer, and fluctuated between 30° and 50° N. Islands had lower BSR than adjacent mainland areas. The NDVI was strongly positively correlated with BSR in mainland areas and on islands. For mainland areas, NDVI explained 65% of the BSR variation, and topographic heterogeneity explained an additional 6% in ordinary least‐squares regression. On islands, NDVI explained 66% of BSR variation, island area explained 13%, and distance to mainland accounted for 1%. Main conclusions In East Asia, we suggest that primary productivity is the key factor underpinning patterns of BSR. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

Correlation between richness per unit area and the species pool cannot be used to demonstrate the species pool effect

Abstract. Significant correlation between the size of the species pool and richness per unit area is usually taken as a demonstration of the effect of the species pool on the species richness per unit area. This is not logically necessary. It has to be emphasized that such a correlation cannot be interpreted as a demonstration of the effect of species pool on the species richness per unit area. More generally, correlation cannot be used to demonstrate a one‐directional effect of one variable upon another, since several types of (functional) relationships could produce correlations between such variables; the same caution has to be taken when analysing the species pool vs. richness per unit area correlation. In this paper, a simple Monte Carlo simulation model is used to demonstrate that (1) statistical processes acting only at the local scale (such as competitive exclusion) can generate exactly the same correlation that has been used to infer the effect of the species pool on richness per unit area; (2) this correlation would be deemed significant even if the mathematical dependency between species pool and richness per unit area is taken into account; and (3) the strength of this relation depends on beta diversity within the contiguous area used to determine the species pool.     

Effects of resource additions on species richness and ANPP in an alpine meadow community

Aims Theories based on resource additions indicate that plant species richness is mainly determined by the number of limiting resources. However, the individual effects of various limiting resources on species richness and aboveground net primary productivity (ANPP) are less well understood. Here, we analyzed potential linkages between additions of limiting resources, species loss and ANPP increase and further explored the underlying mechanisms.Methods Resources (N, P, K and water) were added in a completely randomized block design to alpine meadow plots in the Qinghai-Tibetan Plateau. Plant aboveground biomass, species composition, mean plant height and light availability were measured in each plot. Regression and analysis of variance were used to analyze the responses of these measures to the different resource-addition treatments.Important findings Species richness decreased with increasing number of added limiting resources, suggesting that plant diversity was apparently determined by the number of limiting resources. Nitrogen was the most important limiting resource affecting species richness, whereas P and K alone had negligible effects. The largest reduction in species richness occurred when all three elements were added in combination. Water played a different role compared with the other limiting resources. Species richness increased when water was added to the treatments with N and P or with N, P and K. The decreases in species richness after resource additions were paralleled by increases in ANPP and decreases in light penetration into the plant canopy, suggesting that increased light competition was responsible for the negative effects of resource additions on plant species richness.     

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers’ residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers’ residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m² along a transect and classified the contents by species. We observed 15.5–19.7 species per 0.01 m², which is high richness per 0.01 m² on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.     

Increasing isolation reduces predator:prey species richness ratios in aquatic food webs

The number of species that live in a habitat typically declines as that habitat becomes more isolated. However, the influence of habitat isolation on patterns of food web structure, in particular the ratio of predator to prey species richness, is less well understood. We placed aquatic mesocosms at varying distances from ponds that acted as sources of potential colonists; then we examined how isolation affected the ratio of predator:prey species richness in the communities that assembled. In the final sampling, a total of 21 species (12 prey and 9 predators) of insects, crustaceans, and amphibians had colonized the mesocosms. We found that total species richness, as well as the richness of predators and prey, declined with increasing isolation. However, predator richness declined more rapidly than prey richness with increasing isolation, which lead to decreasing predator:prey ratios. This result conflicts with prior demonstrations of invariant predator:prey ratios in freshwater communities.     

Relative importance of isolation, area and habitat heterogeneity for vascular plant species richness of temporary wetlands in east-German farmland

The purpose of this investigation was to explore the determinants of vascular plant species richness for temporary, isolated wetland habitats which are influenced by hydrologic gradients and characterized by variation in habitat area. The dependent variables total plant species richness and the number of obligate wetland species were analyzed consecutively. In regression analyses habitat area explained between 11 and 15% of the variation in the dependent variables. Habitat area was correlated with the heterogeneity of the hydroperiods between the upper and lower parts of the hydrologic gradients. In multivariate regression analyses, habitat heterogeneity accounted for 70-77% of the variation in the dependent variables, and habitat area did not have a significant impact. The results are most consistent with the habitat heterogeneity hypothesis. I therefore concluded that area is a surrogate variable for habitat heterogeneity which directly enhances vascular plant species diversity. There was no significant impact of isolation on species richness. The data suggest that the expanses of agricultural fields are not an effective barrier to the dispersal of the studied plant species. Only 10 of,52 wetland species were negatively influenced by isolation. This group of species did not differ from the other wetland species with respect to dispersal strategies and longevity of seed banks. However, the longevity of the seed banks was generally high, and there was a dominance of species whose propagules are transported with the soil clinging to the feet of birds. The results are discussed in the context of accurate dispersal strategies and remnant populations, which may counteract the effects of isolation.     

Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns

Summary Habitat subdivision by geography or human activity may be an important determinant of regional species richness. Cumulative species-area relationships for vertebrates, land plants, and insects on island archipelagoes show that collections of small islands generally harbor more species than comparable areas composed of one or a few large islands. The effect of the degree of habitat subdivision in increasing species richness appears to increase with the distance from potential sources of colonists. Mountaintop biotas show no clear differences between species richness on large alpine areas and collections of smaller peaks. National park faunas generally have more species in collections of small parks than in the larger parks. In all cases where a consistent effect of subdivision is observed, the more subdivided collection of islands or isolates contains more species. To the degree that these data provide guidance for establishing nature reserves, they suggest that increasing the numbers of reserves may be an important component of conservation strategies.     

施肥和放牧对青藏高原高寒草甸物种丰富度的影响

在草地生态系统中, 施肥通常会导致生物多样性下降, 但是关于引起生物多样性下降的机制还存在着很大的争议。该研究基于一个4年的施肥实验, 试图通过个体大小的不整齐性和单位植物氮含量, 定性地揭示青藏高原东部高寒草甸施肥后多样性下降的原因。研究显示: 在封育地, 施肥致使个体大小不整齐性增加了15%, 并不同程度地增加了物种的高度。同时, 施肥使物种间单位植物氮含量存在显著差异的数目降低了65%。施肥后光竞争加剧, 导致大个体植物排斥小个体植物, 进而引起了物种丰富度下降29.6%。与封育地不同, 放牧地施肥并没有改变个体大小不整齐性和物种的高度, 而是使物种间单位植物氮含量存在显著差异的数目增加了11.4%。施肥并没有改变放牧地的光竞争强度, 而是增加了物种间对土壤营养元素氮的竞争强度, 进而引起了物种丰富度下降17.3%。该研究还发现, 放牧施肥地的物种丰富度下降速度和等级显著低于封育施肥地的物种丰富度下降速度和等级, 这表明放牧减缓了施肥对物种丰富度的影响力。     

环境因子对太白山高山植被物种组成和丰富度的影响

高山植被是一类具有重要生态和经济价值的植被类型,了解其物种组成和丰富度与环境因子的关系对于该类型植被保护、管理以及植物资源合理开发利用策略的制订具有重要指导意义。基于太白山高山植被和环境因子野外调查及室内实验数据,采用CCA排序法探索了环境因子对物种组成的影响,偏CCA计算了各环境因子对物种组成的总效应和净效应,GLM回归模型拟合了物种丰富度对环境因子的响应。结果表明,13个环境因子共解释了物种组成变异的31.7%,其中海拔、坡度、土壤碱解氮含量、全磷含量、坡向、岩石盖度、p H值、土壤厚度、有机质含量、有效磷含量和全氮含量对物种组成的净效应达显著水平(P0.05),但其作用强度依次减小。GLM拟合结果显示,物种丰富度与环境因子存在4种显著(P0.05)关系,即物种丰富度沿海拔和土壤厚度梯度单调递增,沿坡度和土壤全氮含量梯度单调递减,沿坡向、土壤p H值、碱解氮含量和全磷含量梯度呈单峰分布,与土壤有机质含量和全钾含量呈倒单峰关系。在这些显著的环境因子中,海拔、土壤碱解氮含量,p H值、有机质含量和坡向解释的物种丰富度变异量最大。