In the Wet Chaco region of Argentina, increasing shrub encroachment in savannas over the last few decades has led to important changes in the structure and functioning of the landscape. Some sectors of this territory are characterized by the appearance of circular clusters of woody patches, dispersed throughout the grassland matrix. The increasing size of these patches leads to a gradual change from grassland to dense shrubland. We studied these circular woody patches in the eastern region of the Argentine province of Formosa and characterized the variation in terms of floristic composition, diversity and predominant seed dispersal mode in different size patches. We observed an increase in species richness, diversity and compositional heterogeneity among patches with increasing patch size. Seed dispersal by animals, especially birds, is an important factor in the expansion of these woody vegetation patches within the grassland matrix.

Nomenclature: Zuloaga et al. (1994, 1996a, b). Some native names were included using quotation marks.

Consequences of sulfide toxicity on succession in wet calcareous dune slacks were investigated. Sulfide may exert an inhibitory effect on dune slack plants, but several pioneer species exhibit ROL (Radial Oxygen Loss) and thereby protect themselves against free sulfide. Under oxic conditions free sulfide will be oxiginated to harmless sulfate. However, successive species when not capable of ROL may be sensitive to free sulfide and cannot invade the area. Therefore, the occurrence of free sulfide may have a stabilizing effect on the pioneer vegetation.

Data on the vertical distribution of oxygen, redox and sulfide were collected in mesocosms with Littorella uniflora or Carex nigra, with and without microbial mats and compared to control mesocosms. Also, in situ data were collected in a dune slack on the Frisian Island of Texel.

In the mesocosms, free sulfide was detected only at night-time in C. nigra populated mesocosms and in unvegetated units, but not in L. uniflora vegetated mesocosms. In the field, sulfide and redox profiles showed distinct differences between the groundwater exfiltration and infiltration site of the dune slack. At the exfiltration site, sulfide was only occasionally found; in contrast, measurable amounts of free sulfide were regularly found at the infiltration site of the slack.

Since Phragmites australis dominates in the infiltration site of the slack, the results suggest that free sulfide accelerate the succession, rather than slowing it down by the exclusion of some plant species.

Nomenclature: van der Meijden (1996) for phanerogams; Schaminée et al. (1995) for syntaxa.

Abbreviations: MPN = Most probable number; ROL = Radial oxygen loss.

Maps of plant individuals in (x, y) coordinates (i.e. point patterns) are currently analysed through statistical methods assuming a homogeneous distribution of points, and thus a constant density within the study area. Such an assumption is seldom met at the scale of a field plot whilst delineating less heterogeneous subplots is not always easy or pertinent. In this paper we advocate local tests carried out in quadrats partitioning the plot and having a size objectively determined via a trade-off between squared bias and variance. In each quadrat, the observed pattern of points is tested against complete spatial randomness (CSR) through a classical Monte-Carlo approach and one of the usual statistics. Local tests yield maps of p-values that are amenable to diversified subsequent analyses, such as computation of a variogram or comparison with covariates. Another possibility uses the frequency distribution of p-values to test the whole point pattern against the null hypothesis of an inhomogeneous Poisson process. The method was demonstrated by considering computer-generated inhomogeneous point patterns as well as maps of woody individuals in banded vegetation (tiger bush) in semi-arid West Africa. Local tests proved able to properly depict spatial relationships between neighbours in spite of heterogeneity/clustering at larger scales. The method is also relevant to investigate interaction between density and spatial pattern in the presence of resource gradients.

Abbreviations: CSR = complete spatial randomness; IPP = inhomogeneous Poisson process.

Nomenclature: Hutchinson & Dalziel (1954–1972).

We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Columbia, Canada, and related differences in species composition and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each dominance type varied among climatic regions. The floristic composition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands.

A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we demonstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of compositional variation was shared with other factors. Our results accord with a scenario in which differences in primary environmental factors and humus form properties, the latter accentuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understorey species composition are discussed.

Nomenclature: Qian & Klinka (1998).

Abbreviations: DMB = Drier montane boreal; MMB = Mild montane boreal; WMB = Wetter montane boreal.

In seasonally dry regions of the world fire is a recurring disturbance but little is known of how fire interacts with granite outcrop vegetation. We hypothesize that the floristic composition in granite vegetation, usually attributed to the edaphic environment, may also reflect the impact of disturbances such as fire. Dramatic differences in floristic composition and cover over 13 years and two fires were observed in vegetation on a Western Australian granite outcrop. This was very marked in the first year following the two fires, with annuals and geophytes showing the greatest turnover of species. Even among the perennial shrubs there was considerable turnover in a number of obligate seeders. After the first fire the number of species declined for woody perennials, herbaceous perennials and annuals, remained unchanged for perennial grasses and sedges, and varied with highest richness 4 yr after fire for geophytes. Demographic studies of two endemic woody obligate seeders and three endemic mallee eucalypt resprouters similarly showed dramatic differences within and between species in seedling recruitment following the two fires. Fire does have a significant impact on the floristic composition of semi-arid granite outcrop vegetation communities. Studies on other granite outcrop systems are needed to test the generality of this conclusion.

Nomenclature: follows the database of Western Australian plant names maintained by the Western Australian Herbarium (Paczkowska & Chapman 2000).

Successional patches are a large component of forest ecosystems throughout the world and their vegetation composition is conditioned by multiple factors such as land use history, disturbances, environmental conditions and landscape context. We investigated the relative contribution of historical, environmental, biotic and spatial factors in determining vegetation composition and invasion by exotic species in secondary forest patches of Sierra de San Javier, Tucumán, Argentina. We estimated canopy cover for shrub, vine and tree species distributed over 51 patches with known land use history. We also recorded environmental, historical and spatial variables and used multivariate techniques to explore the relationship between forest composition and explanatory variables. Land use, time since abandonment, altitude, slope and cover of different strata were related to the vegetation pattern in the study site, and they were all significantly structured over space. Exotic species appeared to differ from natives in their response to explanatory variables. Overall, exotic species were dominant on the edges of young patches originated from herbaceous crops, but the total number of exotic species was related to the distance to urban areas and small farms identified as potential sources of exotic propagules. Vegetation composition of secondary forests in NW Argentina was related to historical and environmental factors, but spatial variables strongly influenced vegetation composition as well as the variation in explanatory variables.

Nomenclature: Dimitri (1988); Digilio & Legname (1966).

Abbreviations: GPS = Global positioning system; MRPP = Multi response permutation procedure; NMDS = Non-metric Multidimensional Scaling; PBSSJ = Parque Biológico Sierra de San Javier; UTM = Universal Transverse Mercator system.

This is the first quantitative study of seed bank characteristics in North American alvar habitats. We assessed seed bank density, species richness, and species composition in 75 plots distributed among five alvar sites in Bruce Peninsula National Park, Ontario, Canada, each of which displayed areas of high and low vegetation cover within the alvar and a fully forested perimeter area. Forested habitats immediately adjacent to alvar patches contained minimal seed banks for species restricted to the alvar patches. Open alvars contained less than 1% seeds from woody forest species. This suggests that forest is not invading adjacent alvar habitat via seeds and that adjacent forest does not contain a reservoir of alvar seeds.

When compared to areas on the alvar with high vascular plant cover, areas with low cover contained a slightly smaller viable seed bank, but seed banks from high and low vegetation cover plots had similar species composition and species richness. High vegetation cover plots had slightly higher mean and maximum soil depths compared with low cover plots, but no differences in other physical and chemical parameters. Thus, spatial heterogeneity in plant cover is associated only weakly with heterogeneity in below-ground factors. Despite the availability of seed and soil resources, vegetation dynamics are constrained in areas with low plant cover, and thus alvar community development seems to respond non-linearly to resource availability.

Nomenclature: Newmaster et al. (1998).

Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs' spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high-resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K-function we analysed spatial patterns at three levels: the single-species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand-dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.

Nomenclature: Feinbrun-Dothan & Danin (1991).

Effects of grazing and environment on vegetation structure have been widely acknowledged, but few studies have related both factors. We made 57 floristic samples in a highly variable landscape of mountain grasslands in central Argentina; 26 sample were in fence-lines with contrasting vegetation. For each sample, we recorded topographic and edaphic parameters, as well as grazing intensity indicators. Floristic gradients were analysed with DCA and relations with abiotic and grazing-related variables were detected with DCCA. Floristic axis 1 was explained by edaphic parameters associated to topography, ranging from communities in well drained soils on upper topographic positions to hydromorphic vegetation in poorly drained soils on lower topographic positions. Species richness decreased as soil moisture increased. Floristic axis 2 was associated with present and long-term grazing indicators, and reflected shifts in vegetation physiognomy and species evenness. Tall tussock grasslands, with low species evenness and evidences of low or null grazing intensity were located at one extreme. Tussocks were gradually replaced by short graminoids and forbs towards the centre of the gradient, as grazing increased, and evenness reached a maximum. In degraded sites with heavy long-term grazing intensities, short perennial species were replaced by an annual species, and evenness decreased. The magnitude of changes in floristic composition produced by grazing decreased with increasing soil moisture, and vegetation-environment relationships were stronger in moderate to highly grazed situations than in lightly or non grazed situations.

We examined if the spatial distribution of epiphytes in a forest stand differs between two sexually and three asexually dispersed species. As study species we have chosen two sexually dispersed species, the lichen Collema curtisporum and the bryophyte Orthotrichum speciosum, and three asexually dispersed species, the lichens C. furfuraceum and Leptogium saturninum and the bryophyte O. obtusifolium. All of these species grow on Populus tremula. In P. tremula stands, individual trees may be regarded as distinct patches for the studied species, while the stand represents a ‘landscape” of discrete patches. The study was performed in two relatively homogeneous forest stands. The study revealed that the asexually dispersed species were more common than the sexually dispersed. It was also shown that the sexually dispersed species were more aggregated than P. tremula in both stands. In contrast, the distribution of the asexually dispersed species mirrors, more or less, the distribution of their host, i.e. the spatial pattern shown by P. tremula. It is unlikely that the measured environmental variables strongly influenced the observed spatial distribution. Thus, we suggest that the different dispersal and establishment strategies were important in shaping the local distribution of the species.

Nomenclature:

The decline of species-rich semi-natural calcareous grasslands is a major conservation problem throughout Europe. Maintenance of traditional animal husbandry is often recommended as an important management strategy. However, results that underpin such management recommendations were derived predominantly from lowland studies and may not be easily applicable to high mountain areas. In this study we analyse the importance of traditional low-intensity summer farming (cattle grazing) for vascular plant species diversity of a subalpine region in the northern calcareous Alps in Austria by resampling from an existing dataset on its vegetation. Results indicate a significant long term decline of plant species diversity following abandonment at the landscape scale. In contrast, within-community effects of pasture abandonment on plant species diversity are equivocal and strongly depend on the plant community. We suppose these differences to be due to diet preferences of cattle as well as to the differential importance of competition for structuring the respective communities. From our results we infer that the main mechanism by which pasture abandonment affects vascular plant species diversity, at least during the first ca. 100 yr documented here, are not local-scale competitive exclusion processes within persisting communities. Instead, post-abandonment successional community displacements that cause a landscape scale homogenization of the vegetation cover seem to be primarily responsible for a decline of species diversity. We conclude, that successful management of vascular plant species diversity in subalpine regions of the Northeastern Calcareous Alps will depend on the maintenance of large scale pasture systems with a spatially variable disturbance regime.

Nomenclature: Adler et al. (1994).

Abbreviations: DEM = Digital Elevation Model; GIS = Geographical Information System; WET = Topographic Wetness Index; EROS = Topographic Erosion Index; GLM = Generalized Linear Model; S = Species Richness; H = Shannon Index of diversity; E = Evenness; CC = Number of plant communities; dC = Berger-Parker Index of community diversity.

We studied the germinable soil seed bank of tall-tussock grasslands along an altitudinal gradient in the mountains of central Argentina. We selected 10 sampling plots at three altitudinal levels (1200 m, 1600 m and 2200 m). We assessed the composition of the established vegetation and took ten compound soil samples (0 - 5 cm depth) at each plot in autumn and spring. The soil samples were sieved, chilled, and incubated in a glasshouse to assess the composition of the seed bank. The similarity between the composition of the seed bank flora and that of the established vegetation was low throughout the gradient. Most species did not change their seed bank strategy along the gradient. Seed bank richness and density increased with altitude. Most species had a persistent seed bank at all altitudinal levels, and the proportion of such species increased with altitude. These results suggest that a cold climate directly and/or indirectly favours the formation of seed banks and seed persistence in the soil.

Nomenclature: Zuloaga et al. (1994); Zuloaga & Morrone (1996, 1999).

Plant population biology considers the dynamics of plant modules within stands. However, stands themselves may have considerable regional turnover in space and time. These changes in the number, distribution and size of plant stands generate a dynamic spatial pattern with important implications for the spatial and temporal dynamics of phytophagous insects using these plants as a host. During five successive years we studied the regional distribution and patch dynamics of the creeping thistle Cirsium arvense and the distribution of associated populations of the herbivore Urophora cardui (Diptera: Tephritidae), a specialist stem gall former. The study conducted was in a 15 km² heterogeneous, agricultural area in northeastern Bavaria.

The distribution of the number of plants per patch was skewed with many more small C. arvense patches than large ones. During the five years of study, there was a 50% increase in the number of C. arvense patches, and a decrease in the mean number of plants per patch (= patch size) to less than half the patch size of the first year. Whilst patch size was randomly distributed in space, patch density showed a consistent, non-random spatial pattern. Patch density was spatially autocorrelated, with areas of high or low patch density having a characteristic dimension of ca. 1 km. Patch size was predictable in time and appeared to be regulated by size dependent processes, with the extinction probability of a patch being negatively correlated with its size. Correlated with the decline of C. arvense patch size during the study, the occupancy and total numbers of the herbivore U. cardui had a marked decrease, suggesting that the regional distribution of the stem gall former is not only influenced by patch number but more importantly by the mean patch size. With decreasing patch sizes, U. cardui was faced with an increasingly dynamic landscape due to higher extinction rates of small patches, although the mean distance between host plant patches decreased.

Remaining deciduous forests in the Fennoscandian boreal landscape have high ecological value, and are considered as key components of the forest landscape as well as remnants of a former natural forest type. To improve our understanding of the formation of deciduous forests, we studied past disturbance regimes and vegetation dynamics in three deciduous forests in boreal Sweden using dendro-ecology, pollen analysis and charcoal analysis. We identified three stages in the development of the studied stands. Firstly, the coniferous period (pre 1800), a long-lasting period characterized by frequent fires, livestock grazing and extensive agriculture during which Pinus sylvestris was dominant. Secondly, the transformation period (1800 - 1900), when logging removed most pines from the sites while fire and grazing continued. At the time of the last fire, the sites lacked a local seed source of pines, resulting in a post-fire succession dominated by deciduous species with the capacity to disperse over long distances. Thirdly, the deciduous period (1900 - present), with little or no disturbance from fire, grazing or logging. Thus, the present deciduous stands have their origins in a complex interaction between changes in fire regime, extensive land use patterns and logging, contrary to earlier simplified explanations. We conclude that the complexity of historical patterns of land use, vegetation dynamics and disturbance should be acknowledged in the future when selecting areas for nature conservation and developing models for ecologically oriented forestry.

Nomenclature: Lid (1985).

Whether nitrogen-fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen-fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria-induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria-enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self-replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre-Coriaria stages. These soil changes resulted in three- to sixfold increases in growth of Griselinia when it was grown in Coriaria-enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.

Nomenclature: Allan (1961); Webb et al. (1988); Fife (1995)Edgar & Connor (2000).

Phytosociology is a subdiscipline of plant ecology that describes the co-occurrence of plant species in communities. Gradient analysis and classification are its complementary tools. Various peculiarities and anachronisms of Central European phytosociology conceal its similarity with Anglo-American approaches. Phytosociology deserves to be updated as a part of modern vegetation science that can build on a vast heritage of high-quality data and the tools to store and analyse them in ways that go beyond syntaxonomy. By providing a context to more specialized pure and applied research, it has a crucial role to play in understanding community structure, ecosystem functioning and biological evolution.

Abbreviations: AAPE = Anglo-American plant ecology; CEPS = Central European phytosociology; PS = Phytosociology.

Personal computers of ever-increasing speed have motivated programmers of multivariate software to adapt their programs to be run in Microsoft Windows and Macintosh platforms. Updated versions of these multivariate programs appear more and more frequently and are marketed intensively. In this review we provide a comparative analysis of the most recent versions of three analytical software packages – Canoco for Windows 4.5, PC-ORD version 4 and SYN-TAX 2000. The three packages share two characteristics. First, the most recent versions are now compatible with the most recent Windows platforms and should therefore be accessible for use by virtually all vegetation scientists. Second, they have capabilities for numerous multivariate techniques, although each package has some unique techniques. Thus, any one of the packages will have much to offer the user.

Abbreviations: BC = Bray-Curtis ordination; CA (also, CoA) = Correspondence Analysis; CCA (also, CCoA) = Canonical Correspondence Analysis; CVA = Canonical Variate Analysis, also called Discriminant Analysis; DCA = Detrended Correspondence Analysis; DCCA = Detrended Canonical Correspondence Analysis; MRPP = Multi-response Permutation Procedure; NMDS (also NMS) = Non-metric Multidimensional Scaling; PCA = Principal Components Analysis; PCoA = Principal Coordinates Analysis; RDA = Redundancy Analysis; TWINSPAN = Two-way Indicator Species Analysis.