Small-scale spatial dynamics of plant species in a grassland community over six years

Abstract. In a species-rich mountain grassland in the Krkonosse Mts., Czechoslovakia, data from four permanent plots of 50 cm x 50 cm were recorded annually from 1985 to 1990 to study the spatial dynamics of the species. Plots were divided into 15 x 15 subplots and the number of vegetative units of all plants within each subplot was determined. There was not much net change at the plot level, but the subplots were very dynamic. Two aspects of the spatial dynamics of the species were followed: (1) persistence, i.e. the tendency of the species to remain in the same subplot, and (2) long-distance spreading, i.e. movement to subplots beyond the immediate neighbourhood. Species differed widely in their persistence and longdistance spreading and were classified into mobility types: long-range guerrilla, short-range guerrilla, phalanx and 'sitting’. The mobility types were, to a certain extent, correlated with the growth form of plants, but some species of one growth form showed different types of small-scale dynamics and some species with different growth forms had the same spatial dynamics.     

古田山中亚热带常绿阔叶林群落物种多样性的空间变异特征

古田山常绿阔叶林的群落组成、结构及其维持机制已有许多研究, 但该地区亚热带常绿阔叶林生物多样性空间变异特征还缺乏认识。本文以古田山24 ha大样地(划分为24个1 ha小样地)为基础, 具体分析了α多样性和β多样性在1 ha尺度上的空间变异特征。结果表明: (1)群落第一、二优势物种在各小样地之间变化不大, 但第三优势种变化较大; (2) α多样性变化中, 样地间木本植物个体数量变异最大, 物种丰富度其次, Pielou均匀度指数变异性最小; (3)物种丰富度与植株个体数量、Pielou均匀度指数没有显著的相关性, 与Shannon-Wiener指数呈显著正相关; Shannon-Wiener指数与Pielou均匀度指数呈显著正相关; (4)相邻样地间物种替代速率空间变异较大, 与物种丰富度的空间变化格局有明显差异。这些结果说明尺度对认识群落结构、探讨群落维持机制有重要作用; 由于森林群落是多尺度生态过程作用的结果, 大尺度样地可能有利于更好地揭示森林群落维持机制。     

Small-scale dynamics and species richness in successional alvar plant communities

Species richness and small-scale spatial dynamics (turnover) were compared in four sites during 1990-1993 1) former alvar grassland, now young Pinus sylvestris forest, 2) clear-cut in 1990, 3) clear-cut and grazed by sheep, 4) nearby grassland It was hypothesized that in communities which show large-scale dynamics, small-scale dynamics is also more rapid to support succession Area-based species richness and shoot density increased in the cut and grazed site No significant dynamics was found in shoot-based species richness Therefore the forest was considered to be stable and the cut and grazed site large-scale dynamical (successional) Almost equally high small-scale turnover values were found in all sites, which supports the carousel model for successional communities, but means we had to reject the initial hypothesis The successional and intrinsic small-scale dynamics of the communities studied were two separate processes which acted at different scales, but the scale could not be only spatially determined, it depended also on the shoot density The overgrowing of this former alvar grassland has resulted in the decrease of shoot density and thus the decrease of species richness per unit area, but the total number of species has not changed much Consequently the possibility to restore grassland communities remains     

Dynamics of bryophytes in a chalk grassland

Dynamics of bryophytes were followed in a chalk grassland in 4 plots, 25×25 cm, each subdivided in to 400 subplots, in which every 3–4 months the presence of all bryophyte species was recorded. During the first 14 months no successional trends were visible. However, dynamics on the scale of one subplot were considerable. Degree of dynamics was markedly negatively correlated with original presence in the plots. Several explanations for this correlation are discussed.     

Vegetation change following exclusion of grazing animals in depleted grassland,Central Otago,New Zealand

Abstract. Models of semiarid vegetation dynamics were evaluated to explain changes in the grassland of interior South Island, New Zealand. Annual records were taken for six years of plant species height frequency and percentage ground cover in five plots established in 1986. One subplot at each site was fenced to exclude sheep, one to exclude rabbits and sheep, and one remained unfenced as a control. Records from 1986–1992 were analysed by ordination. The overall pattern of vegetation change shows considerable year-to-year variation. At some sites, variation in vegetation composition between years was as great as, or greater than, that between grazed and ungrazed subplots. Such variation is particularly evident in grazed vegetation, perhaps because it is under greater stress than ungrazed vegetation. At one site changes in vegetation total cover and species composition could be statistically related to rainfall during the first half of the growing season. The only general trends following cessation of grazing were for perennials to increase in frequency, and for year-to-year changes to become smaller with time. Total vegetation cover values seldom changed as a result of cessation of grazing, but tended to follow year-to-year changes in species frequency. The results do not in general support switch/state-and-transition models of semi-arid vegetation dynamics. Vegetation change follows changes in grazing and climate with little lag. This most closely conforms with the Pulse-phase dynamic model.     

High species mobility in species-rich plant communities: An intercontinental comparison

Using results from a long-term study of fine-scale dynamics in grasslands in four widely separated study areas from two continents, we provide further evidence to support the idea of the carousel model as an aid to describe the high fine-scale temporal and spatial species mobility found in grassland communities. Cumulative species numbers on small subplots in plots situated in stable plant communities, determined as the sum of species appearing in these subplots in one or more years over a period of time, are very high. In floristically different species-rich grasslands, varying from moist pine savannas in North Carolina and Mississippi, to humid chalk grassland in the Netherlands and seasonally dry limestone grassland in Sweden, average species numbers on subplots of 0.01 m² in plots of 2.5 m² over the period 1985–1989 were similar, most plots falling in the range 10.8–13.2. The total cumulative species numbers were similar as well, most plots falling in the range 17.4 and 20.9. Yearly average species numbers remained relatively constant. Considerable species turnover is occurring in all these communities; on average three species appear and three disappear each year in each 0.01 m² subplot. Total species accumulation on 0.01 m² subplots over the period 1985–1989 varied considerably, from 4.1 to 11.6, and is correlated with the cumulative species total on the plot, the latter figure being considered as correlated with the size of the species pool.     

Small-scale patch dynamics after disturbance in litter ant communities

The dynamics of re-colonisation of disturbed patches may aid in the understanding of spatial variation of species richness. The present study experimentally tested the hypothesis that the variation of litter ant local species richness and composition is caused by the dynamics of re-colonisation after disturbances. We were particularly interested in whether the re-colonisation was by pre-existent species or species new to the patches, and whether the succession of species evidences the existence of dominance-controlled or founder-controlled communities. Litter patches of a forest remnant in Southeast Brazil were disturbed by removing most animals through litter drying, and litter samples were returned to the same sites from where they were removed. Ant species richness and composition were compared before and 2 months after the disturbance. Dissimilarity among disturbed and non-disturbed samples was compared to infer the succession model occurring after disturbance. Ant species richness did not recover after 2 months, and species composition of the disturbed samples showed more new colonisers than pre-existent species. Dissimilarity among samples in the disturbed plots was smaller than in the control plots, indicating a directional, or dominance-controlled, succession. The changes in species composition observed were caused by a decrease of some species, particularly predators, and an increase of species that are possibly opportunistic. Patches of litter are naturally disturbed in time and space, and evidence from the present paper indicates that succession occurring in these patches would lead to different species richness and compositions. Thus the dynamics of re-colonisation contributes to explaining the diversity of litter-dwelling ant communities at larger spatial and temporal scales. In each patch the succession seems to be directional, with opportunist species re-colonising preferentially empty plots. Therefore, these communities may attain a high diversity due to a small-scale patch dynamics model.     

Small-scale patterns of species richness in Swedish semi-natural grasslands: the effects of community species pools

Temperate semi-natural grasslands are known for their high plant species richness at small spatial scales. We examined the variation in small-scale species richness in a sample of 63 sites from Swedish semi-natural grasslands, located as fragments in the modem landscape dominated by forest and agricultural land. Data were obtained from two spatial scales at each site. 1 dm² and 4 m². Using an analysis based on a Monte Carlo simulation, we found support for the species-pool hypothesis: a high species richness at the I dm- scale was associated with high species richness at the 4 m2 scale. The conclusion from this pattern analysis would, however, be considerably strengthened if we could reduce the likelihood that other mechanisms than sampling from species pools of unequal size influence the pattern of small-scale species richness. Additional analyses were made in order to identify such mechanisms. We examined whether four putative key traits: seed size, seed production, plant size and reproductive allocation were different among species at comparatively species-rich vs species-poor I dm' plots. We found only a little evidence for such differences. There was a weak tendency that species in the plots with high species richness possessed larger (and fewer) seeds than species from species-poor plots. Our results are congruent with the main prediction of the species pool model: variation in small-scale species richness (1 dm²- scale) is basically a result of sampling from unequally sized community species pools (4 m² scale). Variation in species richness between the 4 nr semi-natural grassland "patches" may thus be sought for among mechanisms operating al larger spatial scales than 4 m². We briefly discuss such mechanisms, based on other studies performed in the same study area.     

Patterns of species richness in a limestone grassland under different treatments in relation to spatial scale

Abstract. In an experiment in a limestone grassland on the Baltic island of Öland, SE Sweden, nutrient and water supply, light intensity and grazing regime were altered in 10 combinations during four years in 10 plots of 0.25 m² with subplots of 0.01 m² and 0.0004 m². Only the combined application of fertilizer and shade led to a strong decrease in average species richness (S₁) at all scales. When comparing species numbers summed up over all 10 replicates of each treatment (S_n) at the three quadrat sizes, differences in effect of these treatments were much smaller, and were so already at the finest scale. α-diversity, measured as (S_n - S₁ was quite constant over different scales for most treatments, i.e. diversity did not increase with an increase in scale. The ‘richness ratio’S_n/S₁ decreased with increasing scale, indicating an increasing degree of homogeneity at larger scales. Treatments which only included fertilizer or shade, maintained high species richness; this high richness was also maintained in combination with grazing and could then be explained by the denser packing of vegetation. Patterns of species richness were correlated at the large scale, but not at the finer scales, indicating a high degree of spatial and temporal heterogeneity at the finer scales. With increasing quadrat size species persistence increased which explains the small effect of certain treatments. Clearly, a range of scales has to be sampled in this type of vegetation to be able to measure different patterns, which may occur under different experimental treatments. The finest scale in this study can become too small, when certain treatments result in a coarse-grained vegetation pattern. The quadrat size of 10 cm x 10 cm should be included in the range of scales. It combines accuracy in sampling with efficiency in time effort, a reasonably large number of species sampled, and a strong differentiation in the effects of the various treatments.     

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.     

Detection of plot-level changes in ectomycorrhizal communities across years in an old-growth mixed-conifer forest

Understanding spatial and temporal patterns present in ectomycorrhizal fungal community structure is critical to understanding both the scale and duration of the potential impact these fungi have on the plant community. While recent studies consider the spatial structure of ectomycorrhizal communities, few studies consider how this changes over time. Ectomycorrhizal root biomass and the similarity of community composition were measured at scales up to 20 cm replicated in nine plots and over 3 yr. Soil cores were additionally stratified into three depths. Annual occurrence of the dominant ectomycorrhizal species was constant at larger spatial scales but varied more across years at a fine spatial scale. Turnover of ectomycorrhizal species between years was observed frequently at scales < 20 cm. The ectomycorrhizal community within a plot was more similar across years than it was to other plots sampled in the same year. Our results demonstrate the dynamic nature of the ectomycorrhizal community even in the absence of large-scale disturbances. The potential role of root turnover and drought stress is discussed.     

Small‐scale turnover in a calcareous grassland,its pattern and components

Abstract. We studied small‐scale turnover in a calcareous grassland community in 2 cm × 2 cm subplots within 46 10 cm × 10 cm plots from 1996 to 1999. In different years, 73–84% of the subplots were occupied by rooted plant individuals, but the location of empty subplots varied considerably between years. Seedlings were the most mobile components in subplots – during one year (from July to July), seedlings either appeared or disappeared in on average 85% of all subplots where seedlings had occurred during that year. Within a one‐year period, vegetative ramets appeared or disappeared in an average of 30% of all subplots that had rooted plants at either the beginning or the end of that year. When only previously empty subplots were taken into account, 80% of appearances were due to clonal growth and 20% due to seedling establishment. Ca. 75% of the seedlings, however, established in subplots that already contained a rooted plant individual. There were significantly less generative appearances when a moist summer was followed by a dry one, and significantly more vegetative appearances when a dry summer was followed by a moist one. 24–33% of the established seedlings survived the first year; from the second year onwards the annual survival was greater than 50%. Seedling survival was higher among shortlived species. Generative turnover varied between species, but we found no correlation with life history characteristics or frequency. Vegetative turnover of species was dependent on the year and on species traits. Sedges had a significantly higher turnover than grasses. Species potentially able to move clonally > 10 cm per year had a higher turnover than slower species. Turnover was not dependent on species frequency in the community. Forbs with more seedlings also had a higher vegetative turnover. There were no evident links between spatial mobility and species coexistence; predominating species exhibited contrasting turnover rates, subordinate species had contrasting turnover rates as well. Turnover is important in more general terms as a process resulting in free gaps – arenas for recruitment of whatever species.     

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.     

Divergent small-scale spatial patterns in New?Zealand?s short tussock grasslands

>Spatial studies of ecology rarely look at small-scale spatial community organisation within multiple plots on multiple sites therefore it is difficult to draw conclusions that can be generalised. We hypothesised that small-scale spatial patterns of Festuca tussock grasslands should be consistent within a site and between various sites because their functional ecology is likely to be similar. Tussocks were mapped in 15 plots ranging in size from 56 to 400 m² spread over four sites. Ripley's K, inhomogeneous Ripley?s K and inhomogeneous pair correlation functions were implemented to detect patterns of aggregation, regularity and not significantly different from random at scales up to 300 cm. While Ripley?s K indicated a general trend of regularity up to small scales (c.?20?cm) and aggregation up to larger scales (>40?cm), these patterns were not upheld by the inhomogeneous Ripley?s K and inhomogeneous pair correlation function analyses, which did not yield consistent patterns. Our results suggest that within- and between-site variation of spatial patterns cannot be assumed to be consistent.     

Fine‐scale persistence of boreal bog plants

Abstract. Persistence, or the tendency of a species to remain in its original position without colonizing new sites, is studied for 24 species on the ombrotrophic Northern Kisselbergmossen in SE Norway. Data sets comprise presence/absence in 436 sample plots (16 cm x 16 cm) and 6976 subplots (4 cm x 4 cm) analysed with a 5‐yr interval. Persistence was calculated for both spatial scales, and the observed values were compared with null models of completely random presence/absence of species. Species characteristics (plot occurrences and persistence) were related to depth to the water table and species optima along ecologically interpreted DCA ordination axes. The observed persistence was significantly higher than predicted from the random model for all vascular plants and cryptogams at both spatial scales. All taxonomic groups showed higher persistence at the sample plot scale than at the subplot scale. No general relationship between persistence and depth to the water table was found, but for the cryptogams there was somewhat higher persistence for the less peat‐producing species. The persistence of the vascular plants depended on ramet longevity, growth form and vegetative mobility. In general, the observed persistence of most cryptogams was high, probably because of their perenniality, low growth rates and high reproductive output. Differences in growth‐form and life history, as well as the higher number of occurrences, are the most likely reasons for somewhat higher mean persistence of hepatics and Sphagna than of vascular plants at the subplot scale.     

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.     

Trends and fluctuations in species abundance and richness in Mediterranean annual pastures

Abstract. Floristic data collected from permanent plots during 14 consecutive years are used to model the frequency of the 62 most abundant species in relation to post-ploughing succession, topography and rainfall in annual Mediterranean grasslands located in a Quercus rotundifolia dehesa. The interannual dynamics of species richness are also analysed. From 1980 to 1993, presence/absence data of grassland species were noted in five 20 cm X 20 cm permanent quadrats placed at random in 1980 in 14 permanent plots on a south-facing slope along the topographic gradient. Weekly autumn rainfall data over the 14 years were analysed using a profile attributes index and Hybrid Multidimensional Scaling to arrange the years according to their autumn rainfall pattern. Generalized Linear Models were used to fit the species richness and species frequency according to topographic position, age since the last ploughing episode, total rainfall in the growing season and autumn rainfall pattern using a forward stepwise procedure. The richness model includes all of these variables, and reveals a relatively high goodness-of-fit (71 %). The fact that the meteorological factors play a key role in modelling richness forces us to include them if we wish to use richness as an indicator of the degree of disturbance in these highly fluctuating annual pastures. Models of species dynamics show that although roughly 33 % of the species have a successional behaviour, the majority are more dependent on temporal heterogeneity associated with rainfall or spatial heterogeneity linked to the topographic gradient.     

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.     

Pteridophyte diversity and species composition in four Amazonian rain forests

Abstract. Local variation in individual density, species composition, species richness and species diversity of terrestrial pteridophytes were studied at four sites in the tropical lowland rain forest of western Amazonia. 15 568 pteridophyte individuals representing 40 species were recorded in four plots. The variability among subplots within the same plot was considerable in all the characteristics measured (number of individuals, number of species, species diversity); the square 1‐ha plot was more homogeneous in these respects than any of the three 5 m by 1300 m transects. Species richness was affected by the density of individuals both within and among plots. Density of individuals was not affected by topographical position within any of the plots, whereas in some of the plots both species richness and species diversity were. Clustering and ordination analyses showed that floristically similar subplots could be found in different plots: although there was a tendency for subplots from the same plot to be floristically similar and therefore to group together, many recognized groups included subplots from two or more plots. Both within and among plots, the floristic differences corresponded to topographic position and were probably related to soil drainage. This was also evident in that the abundance patterns of many species followed the topography.     

Dissecting components of population-level variation in seed production and the evolution of masting behavior

Theory predicts that at higher trophic levels species are especially vulnerable to habitat fragmentation due to small population size and dependence on particular prey species. Using aphids as model organism, we tested the hypothesis that herbivore abundance increases in fragmented habitats as a result of reduced predator and parasitoid densities. In a 3 year-study, we examined the population dynamics of aphids with respect to host plant abundance and ant nest density in experimentally fragmented calcareous grasslands at two sites in the northern Jura mountains. Fragments of different size (area: 20.25 m², 2.25 m² and 0.25 m²) were isolated by a 5 m wide strip of frequently mown vegetation and corresponding control plots were situated in the adjacent undisturbed grassland. Aphid density was higher in fragments than in control plots. This was a combined result of a higher frequency of aphid-infested plants and larger aphid colonies in fragments than in control plots. Furthermore, a larger proportion of aphid colonies was ant-attended in fragments than in control plots. Aphid colonies were also more frequently visited by ants in fragments than in control plots in one of the 3 years. Parasitoid pressure on aphids was not influenced by the experimental fragmentation. Neither were aphid species richness and diversity affected by the fragmentation. Our study shows that even small-scale habitat fragmentation can have profound effects on the abundance of herbivorous insects. The effect on aphid density was consistent over 3 years and two sites with slightly different aphid communities.