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.     

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.     

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.     

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.     

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.     

Community relaxation in fragmented landscapes: the relation between species richness, area and age

Estimates of species loss due to habitat destruction are normally based on calculations employing the species–area relation, S = cA ^z . The validity of this approach is based on the assumption that the value of the exponent ( z ) defining the slope of the species–area relation in nonfragmented communities is at a steady state and that z is thus a constant. However, departure from such an assumption renders this approach unreliable. Here I report the results from a natural field experiment using "model" bryophyte-based microlandscapes designed to follow the species richness dynamics of microarthropod communities postfragmentation. Community isolation due to fragmentation initiated a delayed community relaxation process and resulted in substantial local extinction. Over the period of the experiment z declined in the control communities and yet remained fairly stable in the fragmented communities. I conclude that predictions of species loss due to habitat fragmentation that do not take into account the fact that often z may not be a constant may lead to error-prone predictions of future species loss.     

Nutrient enrichment weakens the stabilizing effect of species richness

With global freshwater biodiversity declining at an even faster rate than in the most disturbed terrestrial ecosystems, understanding the effects of changing environmental conditions on relationships between biodiversity and the variability of community and population processes in aquatic ecosystems is of significant interest. Evidence is accumulating that biodiversity loss results in more variable communities; however, the mechanisms underlying this effect have been the subject of considerable debate. We manipulated species richness and nutrients in outdoor aquatic microcosms composed of naturally occurring assemblages of zooplankton and benthic invertebrates to determine how the relationship between species richness and variability might change under different nutrient conditions. Temporal variability of populations and communities decreased with increasing species richness in low nutrient microcosms. In contrast, we found no relationship between species richness and either population or community variability in nutrient enriched microcosms. Of the different mechanisms we investigated (e.g. overyielding, statistical averaging, insurance effects, and the stabilizing effect of species richness on populations) the only one that was consistent with our results was that increases in species richness led to more stable community abundances through the stabilizing effect of species richness on the component populations. While we cannot conclusively determine the mechanism(s) by which species richness stabilized populations, our results suggest that more complete resource-use in the more species-rich low nutrient communities may have dampened population fluctuations.     

Species–area relationships of butterflies in Europe and species richness forecasting

Species–area relationships (SARs) of European butterfly species (Rhopalocera) appear to follow power functions with Mediterranean butterflies having a much higher slope value (z=0.49) compared to the slope for the northern and eastern European countries (z=0.10). A simulated process of species extinction by a stepwise density dependent random elimination of species affected species–area patterns differently. For Mediterranean countries SAR slopes decreased, for other European countries slopes increased during the extinction process. Comparisons of species numbers before and after extinction with those predicted by a classical SAR approach differed widely and revealed that SARs are not able to predict future species numbers at local scales. For Mediterranean countries the classical SAR approach underestimated the number of species remaining after simulated extinction, for all other European countries SARs highly overestimated species numbers. These contrasting patterns indicate that changes in SAR patterns do not unequivocally point to changes in species diversity or community structure as assumed by current theory. On the other hand, the results strongly indicate that simplified applications of SARs for forecasting might give misimpressions about species loss and future biodiversity if the initial community structure, especially relative densities and numbers of species with restricted range size, are not taken into account.     

Elevational gradients in ant species richness: area, geometry, and Rapoport's rule

Studying the distributions of plants and animals along environmental gradients can illuminate the factors governing and maintaining species diversity. There are two general predictions of how species richness and elevation are related: either species richness decreases monotonically with increasing elevation or richness peaks at mid-elevations. Several processes might contribute to this pattern. In this paper, I examine patterns in ant species richness along elevational gradients in three states in the western US: Colorado, Nevada, and Utah. I test for the effects of available area and the geometric constraints model on species richness patterns. I also test Rapoport's rescue hypothesis, which relates the extent of species' elevational ranges to patterns in species richness. In each state, species richness peaked at mid-elevations. Area explained more variation in species richness than the geometric constraints model in Colorado and Utah, but not in Nevada. Area and geometric constraints together explained 90%, 99%, and 57% of the variation in species richness in Colorado, Nevada, and Utah, respectively. Even though there were peaks at mid-elevations, I still found a strong Rapoport effect. This work suggests that the influences of area and geometric constraints cannot be overlooked when examining patterns in species richness along environmental gradients.     

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.     

Indicators of species richness at the local scale in an alpine region: a comparative approach between plant and invertebrate taxa

Studies investigating congruent variations in species richness patterns in alpine habitats are scarce. We investigated the potential of using the indicator taxa approach for species richness in alpine habitats of the Scandes (Norway). In four areas, we investigated seven functional and taxonomic terrestrial groups of organisms and evaluated their contribution to the species diversity. The function of each group as a surrogate for the overall species diversity or for the diversity of another taxon was analysed. Three groups of invertebrates (spiders without Lycosids, Lycosids only, and ground beetles), three groups of plants (shrubs, graminoids, and herbs), and lichens were used for a cross-taxon analysis of species diversity. Congruence in species richness was restricted to several significant results, with vascular plants and spiders (Araneae) being best suited as surrogate taxa in alpine habitats of the Scandes. In the cross-taxon analyses they showed strongest significant positive correlations, covering the total species richness of the alpine habitats best. Species counts in one group account for up to 70% of the variation in total species richness. We found only limited evidence for an ideal, efficient biodiversity indicator taxon that could be applied without restrictions at different alpine habitats in low and middle alpine areas. Thus, our results suggest that it is very important to use more than one taxon as indicator for species richness in terrestrial alpine habitats. This should facilitate future conservation planning in alpine areas.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

Distinguishing area and habitat heterogeneity effects on species richness: Birds in Victorian buloke remnants

Abstract Resolving whether area per se or habitat heterogeneity has the greater influence in controlling species richness remains a controversial yet important question. Here we show that avian species richness of same-sized transects (1 ha) is independent of the remnant area (of buloke woodland) within which a transect is positioned. We also show that avi-faunal similarity of pairs of transects randomly placed within the largest remnants (≥ 48 ha) is not consistently related to either proximity (i. e. being within the same remnant) nor to physiognomic characteristics of the transects. We believe that much of the controversy over area/habitat heterogeneity effects is probably related to scalar issues and propose a protocol by which some resolution of the question might be reached. The protocol involves ‘zoom’ sampling in which successively larger transect sizes are used, and measures of faunal richness and habitat heterogeneity are made at these different grains of resolution. One of our intentions is to stimulate discussion on how heterogeneity might be measured when grains increase from typical transect sizes (ca 1 ha) up to much larger grains (ca 128 ha).     

二道白河河岸带植物群落最小面积与物种丰富度

河岸带是森林小流域单元的重要组成部分之一。由于河水的影响和边缘效应等因素的综合作用,河岸带植物群落与远离河岸带的森林群落在组成,结构和分布格局等方面存在较大差异,其如落最小面积也不同。本文对长白山原始阔叶红松林河岸带植物群落最小面积和物种丰富度进行了探讨。结果表明,河岸带植物群落的最小面积均小于远离河岸带的森林群落的最小面积,在河岸带,阔叶红松林群落的60,80和90％植物在时的平均最小面积分别约为80,180和320m^2;而远离河岸带的森林内部,相应的平均最小面积分别为260,380和480m^2左右。河岸带植物群落的物种丰富度普遍高于森林群落。     

Explaining species richness from continents to communities: the time-for-speciation effect in emydid turtles

Speciation is the process that ultimately generates species richness. However, the time required for speciation to build up diversity in a region is rarely considered as an explanation for patterns of species richness. We explored this "time-for-speciation effect" on patterns of species richness in emydid turtles. Emydids show a striking pattern of high species richness in eastern North America (especially the southeast) and low diversity in other regions. At the continental scale, species richness is positively correlated with the amount of time emydids have been present and speciating in each region, with eastern North America being the ancestral region. Within eastern North America, higher regional species richness in the southeast is associated with smaller geographic range sizes and not greater local species richness in southern communities. We suggest that these patterns of geographic range size variation and local and regional species richness in eastern North America are caused by glaciation, allopatric speciation, and the time-for-speciation effect. We propose that allopatric speciation can simultaneously decrease geographic range size and increase regional diversity without increasing local diversity and that geographic range size can determine the relationship between alpha, beta, and gamma diversity. The time-for-speciation effect may act through a variety of processes at different spatial scales to determine diverse patterns of species richness.     

Small-scale variation in ecosystem CO2 fluxes in an alpine meadow depends on plant biomass and species richness

Characterizing the spatial variation in the CO₂ flux at both large and small scales is essential for precise estimation of an ecosystem’s CO₂ sink strength. However, little is known about small-scale CO₂ flux variations in an ecosystem. We explored these variations in a Kobresia meadow ecosystem on the Qinghai-Tibetan plateau in relation to spatial variability in species composition and biomass. We established 14 points and measured net ecosystem production (NEP), gross primary production (GPP), and ecosystem respiration (Re) in relation to vegetation biomass, species richness, and environmental variables at each point, using an automated chamber system during the 2005 growing season. Mean light-saturated NEP and GPP were 30.3 and 40.5 μmol CO₂ m⁻² s⁻¹ [coefficient of variation (CV), 42.7 and 29.4], respectively. Mean Re at 20°C soil temperature, Re₂₀, was −10.9 μmol CO₂ m⁻² s⁻¹ (CV, 27.3). Re₂₀ was positively correlated with vegetation biomass. GPP_max was positively correlated with species richness, but 2 of the 14 points were outliers. Vegetation biomass was the main determinant of spatial variation of Re, whereas species richness mainly affected that of GPP, probably reflecting the complexity of canopy structure and light partitioning in this small grassland patch.