Direct and indirect effects of nitrogen deposition on species composition change in calcareous grasslands

Atmospheric nitrogen (N) deposition has been identified as a major threat to biodiversity, but field surveys of its effects have rarely focussed on sites which are actively managed to maintain characteristic species. We analysed permanent quadrat data from 106 plots in nature reserves on calcareous grassland sites in the United Kingdom collected during a survey between 1990 and 1993 and compared the data with the results from resurvey of 48 of these plots between 2006 and 2009. N deposition showed no significant spatial association with species richness, species diversity, or the frequency of species adapted to low nutrient conditions in the 1990–1993 dataset. However, temporal analysis showed that N deposition was significantly associated with changes in Shannon diversity and evenness. In plots with high rates of N deposition, there was a decrease in species diversity and evenness, a decline in the frequency of characteristic calcareous grassland species, and a lower number of rare and scarce species. As all sites had active management to maintain a high diversity and characteristic species, our results imply that even focussed management on nature conservation objectives cannot prevent adverse effects of high rates of N deposition. Structural equation modelling was used to compare different causal mechanisms to explain the observed effects. For change in Shannon diversity, direct effects of N deposition were the dominant mechanism and there was an independent effect of change in grazing intensity. In contrast, for change in herb species number, indirect effects on soil acidity, linked to both N and S deposition, were more important than direct effects of N deposition.     

Disturbance,species loss and compensation: Wildfire and grazing effects on the avian community and its food supply in the Serengeti Ecosystem,Tanzania

An important question in biodiversity studies is whether disturbances in ecosystems will cause a net loss of species or whether such losses can be compensated by replacement of other species. We use two natural disturbances, fire and grazing, to examine the response of bird and arthropod communities in grasslands of Serengeti, Tanzania. Both burning and grazing by migrant ungulates take place at the end of the rains in June–July. We documented the communities before disturbance, then 1, 4 and 20 weeks after disturbance on three replicate plots and compared them with three undisturbed plots. Birds were recorded by observation, arthropods from pitfall, tray trap and sweepnet samples. We expected that as the grass biomass was reduced by either disturbance, bird communities would change with concomitant change in arthropod food abundance. Alternatively, bird communities would change not with the absolute amount of food but with the greater accessibility of food as the grass structure changed from long to short grass. Results showed first that both bird species richness and abundance increased after both types of disturbance, but burnt sites showed a greater increase than that for grazed sites. Second, there was a change in bird species composition with disturbance. The functionally equivalent athi short‐toed lark (Calandrella athensis) was replaced by the red‐capped lark (Calandrella cinerea). Third, the abundance of most groups of arthropods was lower on disturbed sites than those on undisturbed sites, and the reduction of arthropod numbers was greatest on burnt sites. These results imply that bird abundance did not occur through an increase in arthropod abundance but rather through a change in the grass structure making food more accessible; and the higher predation could have caused the lower arthropod abundance. In addition, some bird species replaced others thus functionally compensating for their loss.     

Nutrient Limitation in a Fire‐derived,Nitrogen‐rich Hawaiian Grassland1

Grasslands created by grass invasions into shrublands or woodlands followed by fire are now a dominant feature of many seasonally dry environments. In Hawaii Volcanoes National Park, introduced perennial grasses dominate grasslands created by fire in grass‐invaded woodlands. In a previous study, we found that net primary production in these grasslands is substantially lower than in unburned woodlands. Yet, our estimates of annual net nitrogen (N) mineralization showed higher rates in these savannas than in the unburned woodlands, rates that appear to greatly exceed annual N demand by the vegetation. We therefore hypothesized that N should not be limited to the plants growing in these sites. We tested this hypothesis with a 2‐yr fertilization experiment. At peak biomass, we found a 30 percent increase in live biomass in plots with N added and no increase in production with only phosphorus (P) added. N and P together were synergistic, suggesting that co‐limitation or P limitation becomes important when N is more available. Plants responded to added N by increasing individual leaf area and shoot length by over 50 percent. Tissue N was higher with added N; hence, biomass N was substantially higher. Tissue P concentrations declined with N addition but were elevated by P addition despite lack of a growth response to P alone. Overall, N limitation exists despite high annual rates of net N mineralization, and co‐limitation of production by P may occur when N is abundant. Here, asynchrony between plant nutrient demand and N availability may contribute to N limitation.     

Game theory provides no explanation for seed size variation in grasslands

Game theoretical models have been suggested to explain the maintenance of a remarkable variation in seed size across species in most types of vegetation. According to these models, which are based on the existence of a trade-off between seed size and seed number; smaller-seeded species can invade any species mixture due to their numerical advantage, and larger-seeded species can invade any species mixture due to their competitive superiority over smaller-seeded species during recruitment. However, till now, there is very limited evidence for seed size effects on recruitment interactions among different species. An experiment was designed using 16 species in Swedish grasslands, varying 384-fold in seed size. Species were sown pairwise and alone in disturbed versus undisturbed small plots in grazed versus ungrazed grassland, and the resulting recruitment was recorded. Seedling densities in the experimental plots were within the natural range. Both disturbance and grazing had a positive effect on recruitment. Seed size did not affect recruitment except from a tendency that increasing seed size made recruitment less dependent on disturbance. Recruitment of smaller-seeded species was not affected by the presence of larger-seeded species. Larger-seeded species did not generally win in direct contest. These results suggest that game theoretical models do not explain maintenance of seed size variation across species in these grasslands. Alternative explanations for seed size variation are that either small-scale heterogeneity provide conditions favouring a range of different seed sizes or other attributes than seed size effectively determine recruitment.     

Defoliation of Centaurea solstitialis Stimulates Compensatory Growth and Intensifies Negative Effects on Neighbors

Efforts to arrest the spread of invasive weeds with herbivory may be hindered by weak effects of the herbivores or strong compensatory responses of the invaders. We conducted a greenhouse experiment to study the effects of defoliation and soil fungi on competition between the invasive weed Centaurea solstitialis and C. solstitialis and Avena barbata, a naturalized Eurasian annual grass, and Nassella pulchra, a native California bunchgrass. Surprisingly, considering the explosive invasion of grasslands by C. solstitialis, Avena and Nassella were strong competitors and reduced the invader’s biomass by 80.2% and 80.1% over all defoliation and soil fungicide treatments, respectively. However, our experiments were conducted in artificial environments where competition was probably accentuated. When fungicide was applied to the soil, the biomass of C. solstitialis was reduced in all treatment combinations, but reduction in the biomass of the invader had no corollary impact on the grasses. There was no overall effect of defoliation on the final biomass of C. solstitialis as the invader compensated fully for severe clipping. In fact, the directional trend of the clipping effect was +6.4% over all treatments after eight weeks. A significant neighbor × soil fungicide × clipping effect suggested that the compensatory response was the strongest without soil fungicide and when C. solstitialis was alone (+ 19%). Our key finding was that the compensatory response of C. solstitialis in all treatments was associated with an increase in the weed’s negative effects on Nassella and Avena – there was a significant decrease in the total biomass of both grasses and the reproductive biomass of Avena in pots with clipped C. solstitialis. Our results were obtained in controlled conditions that may have been conducive to compensatory growth, but they suggest the existence of mechanisms that may allow C. solstitialis, like other Centaurea species, to resist herbivory.     

Species-Rich Scandinavian Grasslands are Inherently Open to Invasion

Invasion of native habitats by alien or generalist species is recognized worldwide as one of the major causes behind species decline and extinction. One mechanism determining community invasibility, i.e. the susceptibility of a community to invasion, which has been supported by recent experimental studies, is species richness and functional diversity acting as barriers to invasion. We used Scandinavian semi-natural grasslands, exceptionally species-rich at small spatial scales, to examine this mechanism, using three grassland generalists and one alien species as experimental invaders. Removal of two putative functional groups, legumes and dominant non-legume forbs, had no effect on invasibility except a marginally insignificant effect of non-legume forb removal. The amount of removed biomass and original plot species richness had no effect on invasibility. Actually, invasibility was high already in the unmanipulated community, leading us to further examine the relationship between invasion and propagule pressure, i.e. the inflow of seeds into the community. Results from an additional experiment suggested that these species-rich grasslands are effectively open to invasion and that diversity may be immigration driven. Thus, species richness is no barrier to invasion. The high species diversity is probably in itself a result of the community being highly invasible, and species have accumulated at small scales during centuries of grassland management.     

Cotyledon damage at the seedling stage affects growth and flowering potential in mature plants

Seedling herbivory is an important selective filter influencing patterns of plant community composition. Nevertheless, while many of the mechanisms governing seedling selection by herbivores are well established, the effects of tissue loss at the seedling stage on subsequent plant development are poorly understood. Here we examined how the removal of 50 or approximately 100% of cotyledon area from 7-d-old chalk grassland seedlings affected subsequent plant growth and flowering over a 100-d period. Cotyledon damage had a significant effect on growth during the establishment phase for six of the nine species. For two species, significant effects on plant growth were manifest in 100-d-old plants. Of the five species that flowered, three developed fewer inflorescences or flowered later as a consequence of cotyledon damage suffered as a seedling. Our results show that, in addition to the direct effect of herbivory on seedling mortality, more subtle sublethal effects may also influence plant establishment. Reduced growth as a result of cotyledon damage may have implications for plant competition during the establishment phase, and on subsequent reproductive success at maturity.     

Propagule vs. niche limitation: untangling the mechanisms behind plant species' distributions

Distinguishing the roles of propagule limitation and niche requirements in controlling plant species distributions is important for understanding community structure, invasion, and restoration. We used species distribution models based on plant and environmental survey data to assess the strength of species' affinities for particular environmental conditions. We hypothesized that species with statistically detectable environmental requirements were primarily niche-limited, while species with weak habitat affinities were primarily propagule-limited. We tested this hypothesis via a seeding experiment in which we compared species' reproductive fitness in occupied and unoccupied sites. Species that appeared to be niche-limited based on distribution models had lower fitness when planted in unoccupied sites, while species that models suggested were propagule-limited had equivalent fitness when planted in occupied and unoccupied sites. Our results demonstrate that within a single community, both species limited primarily by niche availability or primarily by propagule availability can be identified using observational data.     

A biogeographic delineation of the European Alpine System based on a cluster analysis of Carex curvula-dominated grasslands

Biogeographic delineations within the European temperate mountains remain poorly understood, as there has been little effort to assemble and analyze vegetation relevés covering Pyrenees, Alps, Carpathians and Balkans altogether. Our study tackles this issue by focusing on the widely distributed alpine acidic grasslands dominated by Carex curvula. Cluster analysis of more than 800 vegetation relevés revealed the European-scale spatial patterns of vascular plant diversity in these alpine grasslands. The geographical distribution of floristic clusters was partly congruent with the physiography of European mountains. Southern European ranges (Southern Balkans and Pyrenees) exhibit a high level of endemism and corresponding floristic clusters are well separated from the others. Marked floristic similarities between the Easternmost Alps, the Carpathians, and the Northern Balkans (Stara Planina) supported a major floristic boundary that runs through the Austrian Alps and that is likely the legacy of a shared Quaternary history. Within the Alps, floristic clustering was mainly driven by ecological drivers and not geography. This paper presents the first detailed study of spatial patterns of species distribution within the European Alpine System, based on a comprehensive analysis of within- and between-community species diversity. It shows that the quantitative analysis of large and consistent data sets may question the traditional delineations of biogeographic regions within European mountains.     

Biodiversity effects increase linearly with biotope space

Understanding the influence of environmental variation on the relationship between biodiversity and ecosystem functioning is of theoretical and practical interest. We predicted that the strength of this relationship should increase with available biotope space (the physical space associated with a species’ niche) due to increased niche complementarity between species. In this study, biotope space specifically refers to soil volume which is associated with the niche dimension of nutrient acquisition. We tested our prediction by growing plant communities on a gradient of increasing soil depth and volume, offering increased rooting space to species. Our results provide support for a linear increase of the magnitude of positive biodiversity effects on above‐ and belowground community biomass with increasing biotope space. This increase was caused by complementarity effects between species. Soil erosion may thus reduce intercropping benefits.