Spatial variation in a grassland on soil rich in heavy metals

Abstract. The vegetation in a grassland area in the prefecture of Kilkis (N. Greece), known for its surface Cu-mineralization, was studied. 43 quadrats were established along a transect through an area where the vegetation formed patches of different size. Cover-abundance and frequency estimates for all species were made. Normal Association Analysis revealed five quadrat groups characterized by Trifolium scabrum, Linaria pelisseriana, Anthoxanthum ovatum, Gypsophila muralis and Minuartia hirsuta ssp. falcata. 43 soil samples were analysed for pH, organic C, CaCO₃, total content of Fe, Zn, Pb, Cu, Mn, Ca, Mg, K, Na and soil texture. From a discriminant analysis performed on soil data five soil groups resulted, which are highly related to the five vegetation groups. Thymus sibthorpii, Minuartia hirsuta ssp. falcata and Rumex acetosella are the most important taxa with respect to physiognomy and patchi-ness of the vegetation. The number of species in each group of quadrats is affected by the relative favourableness or severity of the soil conditions. Metal contents, mainly Zn, Cu, Mg and Na, and soil texture are considered to be among the main factors controlling the structure and physiognomy of the vegetation.     

Effects of herbivory on the growth and reproduction of Picris hieracioides in the Mediterranean region

Abstract. The impact of herbivorous insects on the growth and reproduction of Picris hieracioides (Compositae) was followed in three consecutive years by comparing plants treated with an insecticide and untreated plants in an old-field population abandoned for 15 yr (P15). In the third year the interactions between herbivory and competition were analysed as well. Pre-dispersal seed predation of seed heads was investigated in three adjacent old-fields abandoned for one (P1), 15 (P15) and 40 yr (P40) in the region of Montpellier, France. Fields differed in vegetation cover and floristic composition and also in density and above-ground biomass of Picris hieracioides plants (from 246 g per plant in P1, 13.3 g in P15, down to 3.3 g in P40, for the first year of study). The cumulative rainfall between January and July, the period of Picris growth, decreased from 1992 to 1994. Moreover, in 1992, three times as much rainfall occurred in May and June – the period of stem elongation – as in 1993 and 1994. As a consequence, environmental stress increased during the study period. Herbivorous invertebrates had a negative effect on the growth and reproduction of individual plants of Picris hieracioides in P15. The effect was not constant with time, being apparent in only two years out of three. These two years (1992 and 1994) had contrasting climatic characteristics, whereas the year with no apparent effect (1993) was climatically intermediate. When herbivory had effects on growth, the insecticide-treated plants had a significantly higher above-ground biomass and produced twice as many seed heads as the control plants. There was no interaction between herbivory and competition. The percentage of damaged seed heads was significantly lower in the P1-field than in the P40-field. The P15-field had intermediate levels of predation. Predation on seed heads increased whereas biomass and reproductive output decreased between 1992 and 1994 in the three old-fields. Therefore, the results on seed predation (but not for above-ground biomass and reproduction) tend to confirm‘the environmental stress hypothesis’ of Mattson & Haack (1987) that states that plants under stress are more liable to be attacked by grazers.     

Effects of soil erosion on the floristic composition of plant communities on marl in northeast Spain

Abstract. This study explored the validity of three responses of vegetation to increased soil erosion: reduction of vegetation cover, number of species and reduced substitution of species. 201 relevés, including edaphic and geomorphological data, were surveyed in the intensely eroded Eocene marls of the Prepyrenees (NE Spain). Changes in plant species’ presence in relevés from different degradation stages were compared. The level of vegetation degradation was defined as the total phanerogam cover which, in the studied area, was correlated to the degree of soil erosion. The considered trends were validated. Reduction of phanerogam cover and species number were gradual from low to high‐eroded areas. Vegetation degradation explained 48% of the species number variance. In the later stages of degradation a significant substitution of species was not observed, only a lower frequency of occurrence of several species that appeared in the whole set of relevés. Through the process of degradation, 47% of species displayed significantly reduced frequencies as degradation increased, none showed a significant increase in frequency. It is concluded that there are no characteristic species in these plant communities that survive in the severely eroded marls. Among the few species that had increased in frequency, most only colonised favourable micro‐environments.     

Numerical syntaxonomy of the Asplenietalia petrarchae in the Iberian Peninsula

Abstract. A numerical syntaxonomic study of rupicolous vegetation of the Asplenietalia petrarchae in the Iberian Peninsula is described. The study is based on 37 local tables (OSUs) of different rupicolous communities from limestone cliffs found in the eastern half of the Iberian Peninsula. Three classification approaches were followed: (1) ‘hard partition’ techniques, (2) ‘fuzzy partition’ and (3) ordination partition. In order to find a direct solution, we tried an evaluation system based on the inertia associated with a CCA, Canonical Correspondence Analysis, and its relationships with the inertia of a similar CA, Correspondence Analysis. The most effective solutions were found through spatial partitions, and the best partition was obtained with CA considering five segments per axis. Among the fuzzy partitions the best solution was obtained with two clusters and a fuzziness value of 1.20. The best hierarchical partition was Complete Linkage with Chord Distance as a resemblance function. With the GRID procedure applied to the CA diagram, three different groups were distinguished. The first group is related to the Saxifragion boisseriano-reuterani, including five OSUs from Baetic ranges; the second group is very close to the Cosentinio-Lafuenteion rotundifoliae and includes 11 OSUs from thermo-mediterranean areas; the third group, including 21 OSUs basically related to meso-mediterranean areas, is strongly related to the Teucrion buxifolii.     

Above‐ground biomass distribution among species during early old‐field succession

Abstract. The interdependence of species richness and plant biomass has widely been accepted as a general biodiversity rule. However, there is no information about how relationships are established during colonization and how total biomass is distributed among plants. The main objective of this study was to determine the role of several factors which we have hypothetized as affecting biomass distribution among species in an early old‐field community. To test this hypothesis, we conducted an experiment in a deeply ploughed Agropyro‐Rumicion crispi community in the Basque Country (Spain). Light, water and soil nitrogen content were factorially manipulated resulting in eight experimental treatments. We also examined several site features, which could potentially influence final biomass distribution: seed bank composition, soil physico‐chemical heterogeneity and cover and density of the dominant plant species in the community. A partition hypothesis testing approach using Redundancy Analysis was conducted to determine the fraction of biomass distribution variability assigned to each treatment and site feature. The most important species, in terms of biomass, were Agropyron repens, Sinapis arvensis, Arrhenatherum bulbosum and Picris echioides. As a general conclusion, biomass distribution among species during early secondary succession primarily depends on light availability and nitrogen supply. Several soil variables, such as magnesium, calcium and clay contents, also explain a relevant fraction of the biomass distribution among plant species. On the contrary, we found no effect of seed bank composition on biomass distribution. Finally, the total species number and cover of dominant species such as Sinapis arvensis, may determine final biomass distribution.     

The vertical distribution of below-ground biomass in grassland communities in relation to grazing regime and habitat characteristics

Abstract. 40 sites, representing different pasture types in Northwest Spain, were sampled in respect of their floristic composition, distribution of above and below-ground biomass and environmental and physical variables. Five plant community types were identified by classification techniques of plant species composition. These communities were then characterized in terms of the percentage of ground covered by herbaceous and shrub vegetation, stones, rocks and gaps as well as their topographic location and characteristics of the shallow soil (pH, organic matter, nitrogen and calcium content). Bio-mass was assessed in terms of above-ground structures, surface crowns and three below-ground layers to a depth of 10 cm. Three types of grazing regime were distinguished: Concentrated Intense Grazing in early spring (CIG), Extended Intense Grazing throughout the spring (EIG), and Non-Intense Grazing (NIG). Grazing regime showed the highest association with plant community type and three broad categories were identified: xeric stressed pastures, which nevertheless received CIG, mesic pastures with EIG, and three kinds of NIG mesic pastures. The xeric communities had the highest proportion of aboveground biomass, as a consequence of their greater proportion of woody perennials. These xeric communities displayed a more gradual reduction in below-ground biomass with depth than mesic pastures, a likely consequence of the low water content in the upper soil layers. The mesic communities had a high concentration of below-ground biomass in the upper layers when they were intensely grazed. However, when grazing was low (i.e. NIG situations), these communities had greater variability in biomass profiles than any of the other pasture types. Possible causes of the patterns in biomass distribution of the intensely grazed pastures are discussed.     

Restoration of high species density in calcareous grassland: the role of seed rain and soil seed bank

Abstract. A small-scale restoration experiment concerning a species-rich calcareous grassland was conducted over a 7-yr period. 20 yr of abandonment (1971-1990) resulted in both a strong reduction of the number of species and an absolute dominance of the shrub Crataegus monogyna and the grass Brachypodium pinnatum - which contributed 98% of the above-ground herb phytomass. The plot (3 m x 6 m) was cleared in September 1990 and thereafter yearly mown in autumn. Small soil seed bank samples were taken immediately after clearing and again in the autumn of 1996. The inner part of the plot showed a reduced soil seed bank compared to the plot underneath the vegetation outside the plot in 1990. Six years later an increased number of seeds in the soil in this inside area could be recorded mainly in the upper soil layer, although the amount was only 40% compared to the outside or control area. Ca. 20% of the species present in the restored vegetation were recorded in seed traps during the main seed dispersal period in 1997. Restoration of high species richness in the plot was a fast process during the first few years: from seven species in 1990 towards 39 three years later. This was significantly faster than the restoration process of the calcareous grassland at the same slope 20 yr earlier. This was probably due to the increased size of the community species pool at the site as a result of appropriate management during the last decades. Relative species number increase was highest at the largest scale during this experiment, viz. at the total plot area. The success of restoration of species-rich, dry grassland vegetation largely depends on the interaction between several conditions, e.g. seed availability, either originating from the seed rain or soil seed bank, favourable germination conditions and a successful development to a mature, seed producing individual. Germination and establishment strongly depend on suitable vegetation structure as a result of sustainable management.     

Effects of litter accumulation on riparian vegetation: Importance of particle size

Abstract. The floristic effects of river‐borne litter that accumulates in riparian zones may vary in space and time depending on variations in mass and particle size of the deposited litter. To analyse the effects of litter mass and size we applied differentsized litter (natural uncut pieces and powder) to riparian vegetation at different quantities. Vegetation responses were analysed after one season at the community level (total biomass or richness for all species) and species traits (biomass or richness for groups of species). At the community level uncut litter, but not powder, reduced species richness and both uncut and ground litter reduced above‐ground biomass. At the species trait level uncut litter had a stronger effect than powder on species richness and biomass. The only positive effect of litter addition was that powder increased graminoid species richness. The topsoil conditions indicated that the major impact of deposited, river‐borne litter was that it acted as a physical barrier directly preventing established plants from penetrating the litter layer and reducing light and soil temperature.     

Impact of cattle dung deposition on the distribution pattern of plant species in an alvar limestone grassland

Abstract. Seed banks in cattle dung, soil under cattle dung and soil under vegetation and growth response of plant species to the changes in soil nitrogen availability were studied in an alvar limestone grassland on Öland, Sweden, in order to analyse the impact of dung deposition and decomposition on the formation of patches of plant species. Results suggest that patches of four plant species could result from cattle dung deposition and decomposition. Impact of dung could proceed in three ways: (1) by changing the relative abundance of species in the soil seed bank under dung, and/or (2) by influencing the deposition of seeds in the dung, and/or (3) by intensifying the growth of some species through nutrient release. Species patches could result from one or more of these aspects. For instance, patches of Arenaria serpyllifolia may be induced by dung deposition because of the dominance of its seeds in dung, while the pattern of Cerastium semidecandrum and Festuca ovina may be due to the abundance of their seeds in the soil seed bank under dung and their positive growth response to increased nitrogen availability.     

Germination and regeneration mechanisms in Mediterranean degenerate forests

Abstract. Germination and developmental behaviour during juvenile stages are of importance in relation to succession mechanisms acting on plant communities. This is particularly so in the Mediterranean region, where the climatic summer drought causes stress. We investigated and compared germination and development of four tree species in two parts of the French Mediterranean region where the original dominant tree species (Quercus pubescens) has been largely replaced by Quercus ilex in the southern part, and Castanea sativa in the northern part. Observed differences in germination rates and developmental behaviour, especially shootroot ratio and the role of water supply, suggest that other species may eventually replace the current dominant species.     

Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation

Abstract. Four herbaceous plant species of a sand dune area and several herb species of an open early‐successional patch were investigated for the occurrence of a simple relationship between aboveground biomass and plant cover. Without exception linear regressions of aboveground biomass on plant cover were found with slope factors depending on the growth form of the species. These results suggest that (early) growth of herbaceous plants in low and (temporarily) open vegetation is not affected by possible constraints caused by a decreasing ratio of plant cover to aboveground biomass. The obtained linear relationships could be used for rapid non‐destructive determination of aboveground biomass by image‐analysis of cover data.     

Invasibility of a coastal strip in NE Spain by alien plants

Abstract. The alien plant species and the ecological factors that facilitate their invasion to a coastal strip in the Baix Camp region (Tarragona, NE Spain) were studied. A detailed inventory of the area showed that 20% of the plant species, most of them from the American Continent, were aliens, many of which were strongly invasive. At the habitat level, the relationships between the invasive behaviour and a number of autecological, ecological and habitat variables were analysed by means of logistic analyses. In the best model five variables were related to alien plant invasions: climatic affinity, disturbance of the invaded habitat, route of introduction, soil moisture during summer, and life form. To elucidate the environmental factors that could be responsible for alien plant invasions at a regional scale, the number of alien taxa and the environmental features at 13 localities of the western Mediterranean area were analysed. The warm lowlands, subjected to low annual rainfall and high population density, exhibited the greatest reception capacity for alien plants. It is concluded that various specific environmental features – high mean temperatures, the abundance of riparian systems, the existence of biotopes with favourable water balance which harbour a great richness of alien species and the high human pressure to which the enclave is subjected, combined with the ecological requirements of the alien species and certain species attributes (biotype) can interact at different scales and have caused the over‐representation of alien plants.     

Root length/leaf area ratios of chalk grassland perennials and their importance for competitive interactions

Abstract. The use of root / shoot ratios to describe allocation of dry weight to structures for capturing soil resources and light is limited due to other functions of the root and shoot such as support and storage. The ratio of fine-root length to leaf area (RLA) provides a better expression of the relative sizes of above-and below-ground exchange surfaces. Dry matter partitioning, leaf area and root lengths were determined for five species of chalk grassland perennial (Carex flacca, Cirsium acaule, Festuca ovina, Leontodon hispidus and Scabiosa columbaria) by extraction of soil cores from an intact sward. The forb species had a greater proportion of their dry matter below-ground. Interspecific variation in values of RLA was considerable, mean values ranging from 137 m/m² in Cirsium acaule to ca. 27 000 m/m² for Festuca ovina. The implications of this interspecific variation in RLA for the competitive interactions in infertile calcareous grasslands are discussed in relation to phenology and internal nutrient cycling.     

Long-term after effects of fertilization on above-ground phytomass and species diversity in calcareous grassland

Abstract. Long-term after effects on species number and productivity in calcareous grassland were analyzed after cessation of fertilization. Three series of permanent plots were monitored yearly from 1971 to 1993. These series differed in duration of fertilization and in fertilizer composition, notably the amount of nitrogen and phosphorus. Yearly above-ground production decreased in all series after fertilization had been stopped, however at different speeds. The grass/forb ratio also decreased, while species number and Shannon index of diversity (H') increased. In the series where fertilizer treatment was stopped in 1967 (1968-series), productivity and grass/forb ratio decreased equally as in the plots where fertilization with a high nitrogen content was stopped in 1979 (80/Npk-series). However, species number in the plots of the 80/Npk-series increased faster than in the 1968-series. This was probably the result of a higher number of species present in the seed rain from the surrounding vegetation in the period after 1980 than from after 1968. In the plots receiving a high amount of phosphorus (80/nPk series), the productivity decreased more slowly than in the high nitrogen plots (80/Npk-series). At approximately the same total production level, the grass/forb ratio remained higher for another five-year period in the 80/nPk-series. Species number and the Shannon index of diversity increased more slowly in the 80/nPk-series than in the 80/Npk-series. Over a given range of productivity, changes in species number and Shannon index correlated better with the grass/forb ratio than with total above-ground phytomass. Therefore, restoration of species-rich grassland should not only be focused on lowering the yearly production, but also on reducing the grass component of the vegetation.     

How do species dominating in succession differ from others?

Abstract. Data on succession were collected from 15 seres starting on bare ground in man-made habitats (i.e. sites disturbed by various mining activities, bulldozed sites, ruderal urban sites, the exposed bottom of a destroyed water reservoir, and abandoned fields), all in the western part of the Czech Republic, Central Europe. The period for which the data on succession were available ranged from 12 to 60 yr. 56 species reaching dominance in some period of succession were selected (the criterion being: at least 25 % cover in at least one year in any sere); they were compared for biological and ecological traits with other species participating in the seres (167 species with at least 1 % cover in at least one year in any sere), and with the Central European flora as a whole. Significant differences between the species dominating in succession and others were found for the following traits: life form, life strategy, pollination mode, and ability of lateral spread. Dominant species differed from the regional flora in distribution of life strategies, pollination mode and immigration status. The results suggest that an ‘ideal successional dominant’ is a tall, wind-pollinated plant, often a geophyte capable of intensive lateral spread, requiring high nutrient supply and sufficient site moisture. The set of traits contributing to achieving dominance by a species in human-made habitats includes both features occurring independently of phylogenies (life strategy, pollination mode, plant height, moisture demands) and those which dominant species probably share due to their common ancestors (nutrient demands, capability of extensive lateral spread).     

Small-scale plant species turnover in a limestone grassland: the carousel model and some comments on the niche concept

Abstract. This study reports on small-scale changes in the distribution of plant species in a 2.5 m² plot of grazed, species-rich Veronica spicata - Avenula pratensis grassland on shallow, dry, nutrient-poor soil in the Great Alvar area (Stora Alvaret) of southern Öland, southeastern Sweden. Multivari-ate analysis of 0.001 m² and 0.25 m² quadrats within the plot showed that there is little floristic variation without any trend in the plot. Average species richness varied little throughout the study period from 1986 to 1991 with 1986 averages of 7.0 on 0.001m², 16.3 on 0.01 m², and 26.1 on 0.25 m². On 0.001 m²the highest species number found was 12, on 0.01 m², 27. However, cumulative species richness, i.e. species number in the first year plus new species appearing in later years (averaged over 40 quadrats) increased over the same period, on 0.001 m² from 7.0 in 1986 to 14.9 in 1991, and on 0.01 m² from 16.3 to 24.1. Cumulative frequency, i.e. number of quadrats out of 40 where a species occurred in the first year or/and in later years, increased as well; the number of cumulatively highly frequent (> 80%) species increased from 5 in 1986 to 18 in 1991. Species mobility on the scale of the small quadrats used implies both appearance and disappearance from these quadrats. Using six examples, species mobility is shown to vary individualistically, both in rate and degree. We postulate that in homogeneous, grazed, nutrient- and water-deficient environments many species can reach virtually all microsites within the plot, which we express through the idea of the ‘carousel model’. We also question the usefulness of the niche concept and re-interpret it by stating that all species of this plant community have the same habitat niche, while most of them are short-lived and have the same regeneration niche. The essential variation amongst the species is their individual ability to establish or re-establish by making use of favourable conditions appearing in microsites in an unknown, complex spatio-temporal pattern.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

Mediterranean pasture dynamics: the role of germination

Abstract. Meteorological patterns have a decisive influence on the inter-annual dynamics of therophyte pastures under Mediterranean climatic conditions. The germination behaviour of annual pasture species was studied by subjecting two collections of seeds taken from plants and soil-seed banks to two phytotron-simulated weather patterns: early and late autumn rains. Species from these pastures were arranged along a gradient of sensitivity to temperature on the arrival of the first persistent rain. This sensitivity was manifested in both the total germination success of the species and the germination time profile. The different germination patterns of the species can provide competitive advantages depending on the autumn weather conditions.     

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.     

Old-field colonization by Daphne gnidium: seedling distribution and spatial dependence at different scales

Abstract. This paper deals with the spatial distribution pattern of the bird-dispersed plant Daphne gnidium in a 10-yr abandoned field under Mediterranean conditions. Colonization of Mediterranean old-fields by bird-dispersed plants is expected to fit a theoretical model in which (1) seed dispersal follows a negative exponential curve with the distance from the seed source and (2) seedlings are better established under perches (nucleation sensu Yarranton & Morrison 1974). However, post-dispersal processes such as seed predation, seed germination and seedling establishment are also spatial-dependent and can lead to spatial autocorrelation in the seedling distribution within an old-field. Results show that both processes in the model (curve of seed dispersal and nucleation) significantly explained the spatial distribution of the seedlings, but some spatial variance remained unexplained. The semivariogram with the statistical residuals of the model detected spatial dependence at small (< 20 m) and large (> 250 m) distance intervals, indicating that some mechanisms with spatial components, apart from the curve of seed dispersal and nucleation under perches, also determined the distribution of seedlings colonizing fields. At scales below 20-m intervals, semivariance increased indicating that similarity between plots is lost when distance increases between them. This pattern may be explained because the favourable micro-environmental conditions for establishment produced under perches could be extended towards neighbour plots where perches were absent. A flat semi-variogram between 20-m and 250-m intervals shows spatial independence along this range. From 250 m on, the semivariance increased again, indicating spatial dependence at coarse-scale. It is possible that the colonization model failed at this scale because different spatial processes to those included in the model (perch presence and distance to the shrubland) could be controlling seed colonization at coarse-scale.