Recolonization of cleared riverine macrophyte patches: importance of the border effect

Abstract. Recolonization of two experimentally cleared 9 m²-patches by macrophytes in a former channel of the Rhône River, France, was investigated from May to July 1994. Two patterns of recolonization were recognized: (1) Groenlandia densa, Luronium natans and Potamogeton natans invaded the bare areas by propagation from the adjacent vegetation (border effect); (2) Potamogeton pusillus colonized the disturbed areas at random, apparently independently from the position of nearby clumps. An intermediate recolonization pattern was shown by Callitriche platycarpa, Elodea canadensis, Hippuris vulgaris, Ranunculus circinatus and Sparganium emersum. Species recolonization patterns and associated strategies were related to species traits, particularly vegetative and sexual reproduction.     

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).     

Effects of cutting and nitrogen deposition on biodiversity in Cantabrian heathlands

Questions: Does the diversity of heathland vegetation change when subjected to experimental disturbances such as cutting and nitrogen fertilization? Do changes in the vegetation structure negatively affect the regeneration of the dominant species Calluna vulgaris? Is cutting an alternative method of conserving the diversity and maintaining the structure of heathlands in the Cantabrian Mountains? Location: Calluna vulgaris heathlands on the southern slopes of the Cantabrian Mountain range, NW Spain. Methods: A total of 60 plots were treated with different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (56 kg‐N.ha^?1.yr^?1). The changes in the cover values of the species present were monitored over a five year study period. The cover values were used to calculate abundance and species richness. Results: Fertilizing with nitrogen allows biodiversity to increase over time. However, the greatest biodiversity is associated with the cutting plus fertilization treatment, since cutting allows gaps to be opened that are easily colonized by pioneer annual species, while fertilization mainly favours an increase in the mean number of perennial herbs (graminoids and forbs). Increased perennial herb richness also corresponds to a rise in their cover values. The recovery of the dominant woody species in these communities, Calluna vulgaris, is not impeded by the increase in perennial herbs species' cover values. Conclusions: In the Calluna vulgaris heathlands studied, cutting plus fertilization allowed an increase in biodiversity over time. No displacement of the dominant woody species, Calluna vulgaris, is observed due to the presence of the perennial herbs. Cutting patches of heathland is recommended as a mechanism for maintaining high vegetation diversity, when grazing is not possible.     

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 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.     

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.     

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.     

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.     

Establishment of pasture species on cattle dung: the role of endozoochorous seeds

Abstract. The process of colonization of cattle dung patches and its relation to endozoochorous dispersal is analysed in a Mediterranean pasture. Dung pats dropped in spring and winter were marked, and the vegetation developing on them during the first two years of colonization was followed. The influence of endozoochorous seeds on dung colonization is assessed through the study of: (1) the vegetation on spring and winter dung pats and the seed contents of both, which is abundant in the former and negligible in the latter; and (2) the vegetation on spring dung pats compared with that of the surrounding pasture. The origin of plants growing on dung (either seeds in the manure or in the soil seed bank) was traced by studying dung colonization under controlled conditions in pots with manure put on seed-free sand, and on pasture turf. The results indicate that endozoochorous seeds germinate in manure; they are the main source of recovery in gaps generated by dung pats. The micro-succession involved is independent of the type of pasture. A small-scale spatial pattern results in which gaps of old dung are dominated by endozoochorous species. Thus, dung patches enhance the similarity between grazed communities, and the variation within communities.     

Some factors affecting successional change on uranium mine wastes: Insights for ecological restoration

Question: Are growth form and dispersal-mode replacement during vegetation succession in semi-arid Mediterranean conditions affected by the starting quality of the substrate and by site aspect? Location: Central-western Spain. Methods: We monitored successions on three waste materials left after uranium mining: unbroken waste, broken waste and wastes amended with a sandy material (Arkoses); both north and south aspects were also studied on each substrate. Results: The substrate starting quality had the greatest influence on spontaneous succession, separating the poorer quality substrates (broken and unbroken wastes) from the better ones (Arkoses) and two reference communities (Topsoil and Dehesa). The importance of aspect was confirmed then within each substrate type. Most species with a short life span (mostly annuals and a few biennials), together with some woody species on Arkoses, showed no response to age (years following the deposition of new soil). Others short-lived species declined over time on the poorer wastes but not on the better Arkoses. There was a tendency for life form replacement (from thero-phytes to hemicryptophytes) during succession only on the poorer-quality substrates. No dispersal-mode replacement sequence was found. Conclusion: Improving the abiotic conditions of the substrate had a great effect on vegetation succession, but this effect was modified by aspect. Aspect took longer to induce differences in floristic composition on the poorer substrates, where succession was slower. Some trends in species responses to successional change were found by considering species traits, particularly life-form.     

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.     

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.     

Effects of germination inhibition on the dynamics of Quercus ilex stands

Abstract. Germination of Quercus ilex L. in coppice stands of this species in the western Mediterranean Basin was examined and a germination inhibitory process is proposed to explain some germination traits. Germination rate and seedling biomass of Q. ilex were greatly modified by watering acorns with various concentrations of aqueous soil extracts from a Q. ilex coppice stand but also when the acorns were sown in soil from Q. ilex coppice stands. In the aqueous extract experiment, Q. ilex germination and seedling weight both decreased as the aqueous extract concentration increased. In the soil type experiment, Q. ilex soil decreased the Q. ilex germination rate. Comparative studies with Q. pubescens germination (this species, replaced by Q. ilex around 5 000 B.P., is assumed to form the climax vegetation of the region) revealed that Q. pubescens was less sensitive to the aqueous extracts and soils of Q. ilex coppice stands. Inhibition of Q. ilex seed germination could be a major reason for the poor seed regeneration and suggested a possible comeback of Q. pubescens.     

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.     

Effect of local competition on resprouting of Arbutus unedo after clipping

Abstract. To evaluate the effects of local competition on the growth and size variability of sprouts following disturbance in a natural population of Arbutus unedo in Catalonia, plants were cut at the base and a neighbor removal experiment was performed. Removal of neighbors resulted in an increase in the number and biomass of sprouts at 2 and 7.5 months after clipping. Number and biomass of sprouts was also correlated with initial plant size (stump area). Inter-genet competition appeared to be symmetric, and acted to delay the onset of interference among sprouts within a genet. Size variability of sprouts on an individual was positively correlated with their density on the stump, supporting the hypothesis that competition among ramets (sprouts) within a genet is asymmetric. Reduced inter-genet competition from neighbor removal resulted in an increase in the number and biomass of sprouts growing from a stump. This resulted in an increase in the asymmetry of competition among sprouts, and therefore an increase in the size variability of these sprouts.     

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.     

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.     

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.     

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.     

Seed population dynamics on badland slopes in southeastern Spain

Abstract. The dynamics of seed population on slope surfaces were studied to test the hypothesis that the lack of vegetation on badlands is caused by seed removal by erosion. The initial soil seed bank and two years of seed rain and seed removal by erosion were estimated in two small catchments, and a seed balance was constructed. In addition, six rainfall simulation experiments were performed to test the susceptibility of seeds to be removed by overland flow. A variety of soil surface conditions, rainfall characteristics and plot sizes were used in these experiments. Soil seed bank densities are low, but enough for the development of plant cover. Seed losses due to erosion after natural rains were low (< 13 %), and in agreement with seed losses from simulated rainfall experiments. After two years, seed inputs in the seed rain were greater than seed outputs through seed removal, which resulted in a continuous increase in the numbers of seeds in the soil bank. These results point out that seed removal by erosion is not the key factor explaining the lack of vegetation on badlands. It is suggested that other factors, such as those related to seed germination and seedling survival, may play an important role.