Rates of small-scale species mobility in alvar limestone grassland

Abstract. Small-scale species frequency and cumulative species frequency were studied in four plots in limestone grassland of the Veronica spicata-Avenula pratensis association on Stora Alvaret on the Baltic island of Öland, Sweden. Species mobility was expressed as increase in cumulative species frequency in 20 subplots of 100 cm². Observed cumulative frequencies from 1985–1989 in all four plots, and from 1985–1995 in one plot were compared with values following from two null models, a ‘minimal mobility’ model and a random mobility model. In ca. 50 % of the cases the observed cumulative frequency was not significantly different from the random expectation. However, in many such cases the mean annual frequency was either very high or very low. Three ways of calculating the mobility rate are presented though only one is used: (observed cumulative frequency -lowest annual frequency) / expected cumulative frequency. Values × 100 range from 0 to 100. There were slight differences between the four plots which were interpreted in terms of differences in grazing intensity and soil depth. It is stressed that the idea of the Carousel model has never been meant to suggest that all species would show random mobility, which we now quantify, but that species differ in their mobility rate and that the mean rate is much higher than generally realized.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Langfristige Auswirkungen ehemaliger Bewirtschaftungsvarianten auf die Diversit?t von Blütenpflanzen, Moosen und Flechten eines Kalkmagerrasens

Jeschke M., Kiehl K., Pfadenhauer J. and Gigon A. 2008. Long-term effects of former management on the diversity of vascular plants, mosses and lichens in a calcareous grassland. Bot. Helv. 118: 95 – 109. Long-term effects of grassland management on the species richness of vascular plants, mosses and lichens were studied in a calcareous grassland near Schaffhausen (Switzerland). Experimental plots were treated for 22 years (1977 – 1999) with different mowing regimes, annual burning in late winter, or abandonment. From 2000 onwards, all plots were mown annually in autumn. Five years later (2004), we studied the after-effects of the previous treatments by recording the cover of vascular plants, mosses and lichens in nested plots with sizes from 0.01 m² to 16 m². Vascular plant species richness as well as the number of species characteristic for calcareous grasslands were significantly lower in formerly abandoned or burnt plots than in regularly mown plots, with larger differences on small subplots. There was no significant effect of the previous mowing time (July vs. October) or mowing frequency (annual or every two years) on vascular plants. Mosses and lichens had been eliminated by annual burning between 1977 and 1999, and only few moss species re-colonized the formerly burnt plots until 2004. The species richness of mosses did not differ between formerly abandoned and mown plots, but there were more mesophytic and fewer xerophytic species in the formerly abandoned plots. Mesophytic mosses were also more abundant in plots that used to be mown every second year than in those mown annually. Our data indicate that even after five years of similar management, i.e. annual mowing in autumn, the previous long-term management had strong after-effects on species richness and species composition of calcareous grasslands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Eingereicht am 19. M?rz 2008, Angenommen am 30. August 2008, Redaktion: Sabine Güsewell     

Early succession on plots with the upper soil horizon removed

Abstract. Succession was studied on plots with the upper soil horizon removed in an area affected by acidic air pollution in the Kru?né Hory Mts., Czech Republic. 10 permanent 1‐m² plots were marked and vegetation recorded annually using a grid of 100 subplots from 1989 to 1995. Constrained ordination analyses showed that soil texture is the most important environmental factor influencing the course of succession. Its effect on species composition increases with successional age of the plant community. On fine‐grained soils species‐poor communities dominated by grasses (Calamagrostis villosa, Deschampsiaflexuosa) and on coarse‐grained soils species‐rich communities dominated by heather (Calluna vulgaris) developed. Succession proceeded from communities where species composition was determined by diaspore availability towards communities where species composition depended on environmental conditions. Successional communities after 10 yr are more dependent on soil characteristics and consequently environmental determination increases over the course of succession and causes the communities to diverge.     

Invasive alien plants affect grassland ant communities, colony size and foraging behaviour

Ants are dominant members of many terrestrial ecosystems and are regarded as indicators of environmental changes. However, little is known about the effects of invasive alien plants on ant populations, particularly as regards the density, spatial distribution and size of ant colonies, as well as their foraging behaviour. We addressed these questions in a study of grassland ant communities on five grasslands invaded by alien goldenrods (Solidago sp.) and on five non-invaded grasslands without this plant. In each grassland, seven 100 m² plots were selected and the ant colonies counted. Ant species richness and colony density was lower in the plots on the invaded grasslands. Moreover, both of these traits were higher in the plots near the grassland edge and with a higher number of plant species in the grasslands invaded by goldenrods but not in the non-invaded ones. On average, ant colony size was lower on the invaded grasslands than the non-invaded ones. Also, ant workers travelled for longer distances to collect food items in the invaded areas than they did in the non-invaded ones, even after the experimental removal of some ant colonies in order to exclude the effect of higher colony density in the latter. Our results indicate that invasive alien goldenrods have a profound negative effect on grassland ant communities which may lead to a cascade effect on the whole grassland ecosystem through modification of the interactions among species. The invasion diminishes a major index of the fitness of ants, which is a colony’s size, and probably leads to increased foraging effort of workers. This, in turn, may have important consequences for the division of labour and reproductive strategies within ant colonies.     

Sampling-Design Effects on Properties of Species-Area Relationships – A Case Study from Estonian Dry Grassland Communities

Despite widespread use of species-area relationships (SARs), dispute remains over the most representative SAR model. Using data of small-scale SARs of Estonian dry grassland communities, we address three questions: (1) Which model describes these SARs best when known artifacts are excluded? (2) How do deviating sampling procedures (marginal instead of central position of the smaller plots in relation to the largest plot; single values instead of average values; randomly located subplots instead of nested subplots) influence the properties of the SARs? (3) Are those effects likely to bias the selection of the best model? Our general dataset consisted of 16 series of nested-plots (1 cm²–100 m², any-part system), each of which comprised five series of subplots located in the four corners and the centre of the 100-m² plot. Data for the three pairs of compared sampling designs were generated from this dataset by subsampling. Five function types (power, quadratic power, logarithmic, Michaelis-Menten, Lomolino) were fitted with non-linear regression. In some of the communities, we found extremely high species densities (including bryophytes and lichens), namely up to eight species in 1 cm² and up to 140 species in 100 m², which appear to be the highest documented values on these scales. For SARs constructed from nested-plot average-value data, the regular power function generally was the best model, closely followed by the quadratic power function, while the logarithmic and Michaelis-Menten functions performed poorly throughout. However, the relative fit of the latter two models increased significantly relative to the respective best model when the single-value or random-sampling method was applied, however, the power function normally remained far superior. These results confirm the hypothesis that both single-value and random-sampling approaches cause artifacts by increasing stochasticity in the data, which can lead to the selection of inappropriate models.     

Small-scale plant mobility in a species-rich grassland

Abstract. Plant mobility was studied in a species-rich grassland in S. Moravia (Czech Republic) at scales from 0.0025 to 2.25 m². Cumulative species numbers, cumulative species frequencies and the distribution of distances between sites of occurrence in particular years were established using data collected from 1991 to 1997. The observed values were compared with null models of completely random and restricted random movement of the plants. Most plants persisted at a spot more frequently than expected from the completely random model and with a few exceptions they also spread to neighbouring subplots more often than expected. Cumulative species numbers were between the ranges predicted by the two random models and increased linearly during the 7-yr period at all scales, indicating a large species pool. The role of clonal spreading and of generative reproduction depended on the growth form of the species. I conclude that a high species richness is not necessarily linked with a high plant mobility, even if coexistence of plant species may be promoted by plant mobility.     

Disturbance,drought and dynamics of desert dune grassland,South Africa

A seven-year study of marked plants and plots in Stipagrostis ciliata (Desf.) de Winter dune grassland, in the arid (<100 mm yr^–1) Bushmanland area of the Northern Cape province of South Africa, was designed to test the hypothesis that establishment of ephemeral plants, and recruitment of perennial grasses was dependent upon disturbances that reduced the density of living perennial grass tussocks. In 1989, eight 4 m² plots were cleared of perennial vegetation by uprooting and removing all plants so as to resemble small-scale disturbances made by burrowing mammals or territorial antelope. The vegetation on the cleared plots and surroundings was monitored until 1996. Initial results supported our hypothesis. In wet years, when ephemeral plants were abundant, their average fresh mass was 2–3 times greater per unit area on the cleared plots than in control plots in adjacent, undisturbed grassland. Many Stipagrostis seedlings established in the cleared plots over the two years following clearing but were rare in adjacent areas among established conspecifics. However, a drought in 1992 (11 mm of rain over 12 months) lead to widespread mortality of the perennial grass, killing 56% (range 22–79%) of established tufts. High densities of Stipagrostis seedlings appeared following the drought-breaking rains in January 1993, both in the disturbed plots and in the surrounding `undisturbed' dune grassland. Ephemeral plants established in large numbers throughout the area during the high rainfall year of 1996 and were generally more numerous in the old disturbances than in control plots. Seven years after clearing the biomass of grass on the cleared plots was approximately 34% of the mass removed from the plots in 1989 whereas in the undisturbed grassland biomass was 66% of 1989 levels. Drought had little long-term effect on community composition, and Stipagrostis ciliata constituted 94–98% of plant community before and after drought. Cleared plots were recolonised by S. ciliata, but the contribution of other grass species increased by 6–9%. Synchronous recruitment following occasional drought-induced mortality can generate even-aged populations of the dominant desert dune grasses.     

Size and heterogeneity rather than landscape context determine plant species richness in semi‐natural grasslands

Question: Is plant diversity in fragmented semi‐natural grasslands related to present and historical landscape context? Location: Southern Sweden. Methods: Plant diversity was described at 30 semi‐natural grassland sites in terms of total and specialist plant species richness at the site and species density at different scales (0.5–10 m²). These measures are commonly used to assess conservation value of semi‐natural grasslands. Landscape context was measured as contemporary connectivity to other semi‐natural grasslands, historical connectivity 50 years ago, amount of linear elements potentially suitable for dispersal (road verges, power line clearings), and amount of forest (inverse of the openness of the landscape). Results: The diversity measures were generally correlated with each other, implying that species richness in a subset of the grassland can predict the total richness. Plant species density at three scales (0.5 m², 10 m² and total) was related to the landscape context using an information theoretic approach. Results showed that total species richness increased with increased size of grasslands, contrary to earlier diversity studies in semi‐natural grasslands. Larger grasslands were more heterogeneous than smaller grasslands, and this is a likely reason for the species‐area relationship. Heterogeneity was also of high importance at the smaller scales (0.5 m², 10 m²). With increased amount of forest, total species richness increased but species density on 10 m² decreased. There was no influence of connectivity in either the contemporary or the historical landscape, contrary to previous studies. Conclusions: Grassland size and heterogeneity are of greater importance for plant diversity in semi‐natural grassland, than grassland connectivity in the landscape.     

High Plant Diversity of Grasslands in a Landscape Context: A Comparison of Contrasting Regions in Central Europe

Some regions and habitats harbour high numbers of plant species at a fine scale. A remarkable example is the grasslands of the White Carpathian Mountains (Czech Republic), which holds world records in local species richness; however, the causes are still poorly understood. To explore the landscape context of this phenomenon and its relationships to diversity patterns at larger scales, we compared diversity patterns in grasslands and other vegetation types in the White Carpathians with those in nearby regions lacking extremely species-rich grasslands, using data from vegetation plots and flora grid mapping of entire landscapes. Although small-scale species richness of grasslands and ruderal/weed vegetation of the White Carpathians was higher than in the nearby regions, the number of grassland and ruderal/weed species in the regional flora of the White Carpathians was not. Diversity of forests was not higher in this region at any scale. Thus the remarkably high local species richness of the White Carpathian grasslands does not result from a larger grassland species pool in the region, but from the fine-scale co-occurrence of many grassland species in this landscape, which results in the formation of grassland communities that are locally rich but with similar species composition when comparing different sites (i.e. high alpha but low beta diversity). This pattern can be partly attributed to the large total area of these grasslands, which reduces random extinctions of rare species, low geological diversity, which enables many species to occur at many sites across the landscape, and high land-cover diversity, which supports mixing of species from different vegetation types.     

Plot shape effects on plant species diversity measurements

Abstract. Question: Do rectangular sample plots record more plant species than square plots as suggested by both empirical and theoretical studies? Location: Grasslands, shrublands and forests in the Mediterranean‐climate region of California, USA. Methods: We compared three 0.1‐ha sampling designs that differed in the shape and dispersion of 1‐m² and 100‐m² nested subplots. We duplicated an earlier study that compared the Whittaker sample design, which had square clustered subplots, with the modified Whittaker design, which had dispersed rectangular subplots. To sort out effects of dispersion from shape we used a third design that overlaid square subplots on the modified Whittaker design. Also, using data from published studies we extracted species richness values for 400‐m² subplots that were either square or 1:4 rectangles partially overlaid on each other from desert scrub in high and low rainfall years, chaparral, sage scrub, oak savanna and coniferous forests with and without fire. Results: We found that earlier empirical reports of more than 30% greater richness with rectangles were due to the confusion of shape effects with spatial effects, coupled with the use of cumulative number of species as the metric for comparison. Average species richness was not significantly different between square and 1:4 rectangular sample plots at either 1‐ or 100‐m². Pairwise comparisons showed no significant difference between square and rectangular samples in all but one vegetation type, and that one exhibited significantly greater richness with squares. Our three intensive study sites appear to exhibit some level of self‐similarity at the scale of 400 m², but, contrary to theoretical expectations, we could not detect plot shape effects on species richness at this scale. Conclusions: At the 0.1‐ha scale or lower there is no evidence that plot shape has predictable effects on number of species recorded from sample plots. We hypothesize that for the mediterranean‐climate vegetation types studied here, the primary reason that 1:4 rectangles do not sample greater species richness than squares is because species turnover varies along complex environmental gradients that are both parallel and perpendicular to the long axis of rectangular plots. Reports in the literature of much greater species richness recorded for highly elongated rectangular strips than for squares of the same area are not likely to be fair comparisons because of the dramatically different periphery/area ratio, which includes a much greater proportion of species that are using both above and below‐ground niche space outside the sample area.     

Seed Bank Effects on Recovery after Disturbance in Reconstructed Tallgrass Prairies

Litter‐removing disturbances such as fire in grasslands temporarily increase available resources for plants, opening a window of opportunity for new establishment as communities recover. At this time, new individuals or species could be added to the community as a result of germination from the local seed bank. In reconstructed grasslands this may be problematic, as the seed bank may contain a suite of undesired species reflective of prior and surrounding land uses. In two, 25‐year‐old, low‐diversity reconstructed grasslands, we tested the effect of local seed bank establishment following litter‐removing disturbance using seedling removal plots (1 m²) and plots where natural seedling establishment was allowed. Following disturbance, the vegetation was either left intact or hayed to enhance seedling establishment (a common practice following inter‐seeding efforts). Although the seed bank and seedling community were dominated by resident grasses (Andropogon gerardii and Poa pratensis), recruitment from the seed bank increased species richness and reduced evenness through the addition of forb species (including Cirsium arvense) in one of the study sites. Haying temporarily altered the abundances of the dominant grasses, but did not consistently affect seedling recruitment. Disturbances that facilitate seed bank recruitment may promote establishment of undesired species within reconstructed grassland communities, and we need to take steps to better manage the contributions into and recruitment from the seed bank to reconstruct sustainable grasslands.     

Model estimation of energy flow in North American grassland bird communities

Summary The energy demands and general food consumption rates of bird populations breeding in North American grasslands are estimated using a simulation model which employs information on population natural history and individual metabolism gathered from several study locations. The total breeding season energy demand of the grassland/shrub-steppe avifaunas ranged from 0.89 kcal m^-2 season^-1 in arid shrub-steppe to 2.92 kcal m^-2 season^-1 in a mesic tallgrass prairie. There was substantial variation between years and between census plots, however, and in general the average avian community energy demands did not differ significantly over the range of locations. Production accounted for 0.9 to 1.5% of the total seasonal energy demand. Roughly 11 to 18% of the seasonal energy flow was required in the production of eggs and maintenance and growth of nestlings and fledglings.On the average, between 209 and 386 kg dry wt km^-2 of prey were consumed by the bird communities breeding in the grassland locations. Seeds contributed more to the total biomass consumed at the drier plots, but in general, animal prey types comprised roughly 80% of the total biomass eaten. Phytophagous insects were the major component of the animal prey.These low magnitudes of energy flow and biomass consumption attest to the relatively minor role of birds in the processing of energy and biomass in grassland ecosystems. If these populations do play an importnat role in the functioning of grassland ecosystems, it must be quite subtle and indirect.     

Changes in chalk-grassland structure and species richness resulting from selective nutrient additions

Abstract. A series of fertilization experiments was carried out over a 5-yr period in a chalk grassland in Limburg (The Netherlands) as part of a study of the maintenance of species richness in species-rich grasslands. Phosphorus and nitrogen were shown to be the most limiting nutrients. Addition of both elements doubled above-ground production, and species richness dropped ca. 50 % in 0.01-m² subplots, relative to controls. However, neither the above-ground production nor plant growth-forms were sufficient to explain the observed changes in species richness. Small-scale structural heterogeneity of the vegetation is probably critical for maintaining high levels of richness. Historically, high nitrogen, low phosphorus conditions were rarely encountered in the Dutch landscape and few species appear adapted to these conditions. Among the chalk grassland species, Brachypodiumpinnatum seems well adapted to these conditions, where it dominates and excludes most other species. A detailed understanding of the small-scale processes responsible for maintenance of species richness is critically important in efforts to maintain the biodiversity of natural ecosystems.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Species richness and vegetation structure in a limestone grassland after 15 years management with six biomass removal regimes

Species richness, species composition and standing crop were studied in a limestone grassland in northern Switzerland after 13 and 15 years of experimental management with 6 management regimes differing in form, time and frequency of biomass removal. The management regimes with a cut at least every 2nd year had a higher species number per 40 m² and 1 m² than those cut less frequently, abandoned or annually burned. Annual burning and a cut every 5th year lead to a similar decrease in species richness as abandonment. Differences in species richness between different regimes increased with increasing spatial resolution of observation. Many species occurred in a plot during only one of the two sample years. This turnover of species was the highest in plots with the lowest species numbers, and increased with increasing spatial resolution. Standing crop was highest in plots burned annually (290 g m^?2), and lowest in plots cut annually in July (1989 g m^?2). Standing crop and litter increased with decreasing frequency of cutting. *** DIRECT SUPPORT *** A02DO006 00005     

Response of Grassland Birds in Sand Shinnery Oak Communities Restored Using Tebuthiuron and Grazing in Eastern New Mexico

Sand shinnery oak (Quercus havardii) communities are a unique component of grassland bird habitat in eastern New Mexico and have been impacted by human activities for decades. These communities are frequently managed with livestock grazing and herbicide application for shrub control, strategies that potentially can be used to restore the historical shrub–grass composition of this plant community. During spring migration and the breeding seasons of 2004 and 2005, we compared density and community structure of grassland bird species among four combinations of tebuthiuron application and grazing treatments that were being evaluated for restoration of shinnery oak communities. We performed biweekly point transects on sixteen 65‐ha study plots in these communities. Density of all avian species combined did not differ between grazed and ungrazed plots. Tebuthiuron‐treated plots had a 40% higher average density for combined species than untreated plots. There was a 41% higher average density of all species during spring 2005 than 2004, but density was similar during the breeding season of both years. These trends were predominantly influenced by densities of migratory Cassin’s Sparrow (Aimophila cassinii), which were greater in tebuthiuron‐treated plots in both years. Densities of resident Meadowlarks (Sturnella spp.) exhibited little response to tebuthiuron or grazing treatments. Avian species richness, evenness, and diversity were only minimally affected by the tebuthiuron and grazing treatments. This study occurred over a period of highly variable precipitation, so future assessments, spanning longer wet–dry cycles and maturing plant communities, may be necessary to completely determine avian response to these restoration efforts.     

Positive diversity–invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale

Background and Aims

Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community.

Methods

In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm² quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm²) within plots (800 cm²) and between plots within the site (approx. 400 m²) over 46 months.

Key Results

Invader success was positively related to resident species diversity and richness over a range of 28–37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders.

Conclusions

Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity–invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.     

Multi-scale analysis of plant species richness in Serengeti grasslands

Aim To assess scale dependence between environmental factors and plant species richness. Additionally, we aimed to identify the scales at which niche relations and habitat heterogeneity, as hypothesized by A. Shmida & M.V. Wilson (1985) Journal of Biogeography, 12 , 1–20, operate in the savanna grasslands that were the focus of this study. Location Savanna grassland plant communities of Serengeti National Park, Tanzania. Methods Plant species richness was sampled in 102 modified Whittaker plots and tested for associations with two climate factors, mean annual rainfall (MAP) and potential evapotranspiration (PET), and two landscape variables, plot aspect (ASP) and topographic variation (TOPO), using multiple regressions. Scale dependence was assessed by conducting regressions after altering three aspects of spatial scale: grain, extent and focus. Grain was altered by analysing plant richness at 1, 10, 10² and 10³ m²; extent was investigated by restricting the maximum distance between samples to 75, 100, 125 and 150 km; and focus was manipulated by averaging samples spatially according to geographical land regions. Within the context of our data, we assumed that niche relations were represented by climate factors and habitat heterogeneity by landscape factors. Results Across all 102 plots, plant species richness between 1 and 10² m² had a negative relation to PET and a weak positive relation to MAP. Plant species richness at 10³ m² had a positive association with TOPO and weaker associations with climate factors. ASP stayed in the model between grains of 10 and 10³ m², but had a very weak positive association with richness. When the focus was changed to land regions, associations between plant species richness and explanatory variables strengthened, but were not qualitatively different. At spatial extents of 75 and 100 km, PET was the strongest correlate of plant species richness across all spatial grains. At spatial extents ≥ 125 km, PET explained the majority of the model variance at spatial grains ≤ 10² m², whereas TOPO explained equal amounts or more of the model variance at spatial grains of 10³ m². Main conclusions Both climate and topographic variation explained plant species richness in Serengeti grasslands, but specific patterns depended on grain, extent and, to a lesser degree, focus. Consistent with the ideas of Shmida & Wilson (1985) , determinants of plant species richness shifted from niche relations to habitat heterogeneity between spatial grains of 1 and 10³ m², although this occurred only at relatively large spatial extents (≥ 150 km). Finally, the signs, strength and shape of plant species richness relationships in Serengeti closely match those that describe macro‐scale patterns of woody plant species richness across the entire African continent.