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.     

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.     

Climatic discrimination of Mediterranean broad-leaved sclerophyllous and deciduous forests in central Spain

Abstract. Climatic differences between three types of deciduous (Quercus pyrenaica) and three types of sclerophyllous (Quercus rotundifolia) Mediterranean forests in the Spanish Sistema Central were analyzed by means of Canonical Discriminant Analysis and Jancey's Discriminant Analysis, applied in successive steps to data from 252 meteorological stations. Climatic data included temperature and precipitation records as well as bioclimatic indices. Discriminant analysis was applied to broad-leaved sclerophyllous and deciduous forest communities sampled at each meteorological station using phytosociological methods. Annual and seasonal (summer, spring) water availability are the most important factor controlling the distribution of the two physiognomic forest types; southwestern associations of Quercus pyrenaica and Q. rotundifolia differ from their colder homologues by annual and monthly temperatures; western associations were separated from eastern ones in terms of annual and seasonal precipitation gradients. Discriminant analysis was a good technique to explore climatic gradients not shown by other general ordination or classification methods.     

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.     

Effect of time on the natural regeneration of salt marsh

Abstract. The effect of time on natural regeneration of two salt marshes was studied in relation to plant and edaphic factors. The study was carried out in two naturally restoring salt marshes, differing in restoration time, in Txingudi (Bay of Biscay). After 20 yr, the younger salt marsh had the same plant species richness and high species similarity as a 35 yr old salt marsh (17 and 16, respectively, similarity index = 0.9), but both sites had lower species richness and similarity than a nearby natural salt marsh (36 plant species and similarity indices of 0.45 with the 35 yr old marsh and 0.46 with the 20 yr old marsh). Plant species present in the two recovering salt marshes followed a similar distribution pattern in relation to organic matter, conductivity and moisture content although this zonation differed from the natural salt marsh. The range of edaphic factors measured was also similar, but differed from those in the natural salt marsh. The process of plant species recolonization and spatial distribution might be delayed by a low probability of species arrival and by the time need for the restoration of hydrologic and edaphic factors. This study supports the necessity of long‐term monitoring in measuring coastal salt marsh restoration.     

The role of spatio‐temporal heterogeneity in the establishment and maintenance of Suaeda maritima in salt marshes

Abstract. The effects of disturbance and microtopography on the organization and dynamics of plant communities were studied in a European salt marsh located in the Bay of Mont St. Michel, France. The existence of seed trapping mechanisms was also tested. The study took place in the lower and middle marsh plant communities dominated by the perennials Puccinellia maritima and Halimione portulacoides, respectively and associated with the annual Suaeda maritima. Three treatments were used in series of plots placed in each community: (1) vegetation removal and root destruction to a depth of 10 cm and refilling, (2) non‐remnant herbicide treatment without vegetation removal and (3) creation of depressions (20 cm deep). These treatments were compared with adjacent control plots. The first year of the experiment showed that the perennials facilitated the establishment of Suaeda by trapping its seeds. Estimation of cover, density and biomass over 5 yr following the disturbances showed that in the first 2 yr Suaeda dominated the disturbed plots. Thereafter Suaeda was gradually eliminated by competitive exclusion after ca. 3 yr in the zone originally dominated by Puccinellia maritima and after 4 yr in the zone occupied by Halimione portulacoides. Depressions constituted refuge habitats for Suaeda by limiting competition with the perennials but also led to a high risk of mortality with temporal fluctuations in density. Despite a period of investigation limited to 5 yr, our study demonstrated that natural disturbances of various types occurred and influenced the dynamics of Suaeda, Halimione and Puccinellia. We deduced that natural disturbances and microtopography are responsible for the maintenance of the habitat in a state of non‐equilibrium by favouring the establishment of both spatial and temporal environmental heterogeneity. These conditions appear to be particularly favourable for the maintenance of annual species such as Suaeda maritima.     

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.     

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.     

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.     

Management of a semi‐natural grassland; results from a 15‐year‐old experiment in southern Sweden

Abstract. Vegetation changes in a semi‐natural grassland of wooded meadow type that had been grazed for centuries are described following the introduction of various management regimes: mowing each year, mowing every third year, burning, mechanical removal of woody plants, chemical treatment of woody plants, continuous grazing and abandonment. The experiment was established in southern Sweden in 1972 and has been in progress for 15 years. In 1972, 1980 and 1986 the botanical composition in these plots was investigated in permanent subplots. The study clearly demonstrates that mowing or grazing is necessary to preserve community structure and that mowing is to be preferred in cases where maintaining species richness is of primary concern. Mowing every third year delayed vegetation change and prevented woody species from spreading. Therefore, periodic mowing might be an alternative way to preserve the flora. In contrast, yearly burning does not seem to be a viable management in this type of semi‐natural grassland. To preserve the open landscape regeneration of woody plants has to be prevented. However, in plots where woody plants were removed the typical grassland flora declined. Abandonment resulted in closed forest.     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Effects of disturbance caused by traditional Spanish rural land use on the regeneration of Cytisus multiflorus

Abstract. Cytisus multiflorus is a leguminous matorral shrub native to the NW Iberian Peninsula, where it is one of the most important species in the extension of matorral at the expense of set-aside agricultural land. Dehesas have traditionally been used for extensive livestock raising and matorral was periodically burnt, cut or pulled out. The two latter practices are now out of use. However, burning is more frequent than in the past. The effects of fire, cutting, and pulling out of C. multiflorus on its regeneration was studied in order to ascertain whether the presently increasing distribution of the species relates to fire-stimulated regeneration or to the reduction of other traditional practices. Three years after treatment two sets of parameters were determined: 1. Plant origin: seedlings and different ramet types (ramets=resprout clumps), density, weight, and biomass as well as the percentage of resprouting. 2. Seed persistence at various soil depths. The possible mechanisms of breaking dormancy and plant emergence in different years after fire were studied in other experiments. The results suggest that the regeneration mechanism in C. multiflorus is stimulated by fire, but it is not an exclusive relationship. Stimulation occurs also after other disturbances leading to the total elimination of aerial biomass. The present expansion of the species does not appear to result from the abandonment of some traditional practices, such as cutting or pulling out, but from frequent fires (resulting in aerial-biomass elimination).     

Chemical and morphological variation of Mediterranean woody evergreen species: Do plants respond to ungulate browsing?

Abstract. Plant defense theories suggest that chemical or structural defences should be maximized when and where browsing is most likely to occur. We tested this hypothesis on four evergreen woody species growing in a Mediterranean area with a high density of ungulates. In this system, levels of browsing are more intense in the winter (due to the lack of annual plants) and young foliage is often preferred. Therefore we predicted that the chemical defences of these species, namely their phenolic content, would vary with leaf age, season and damage intensity. In addition, we tested whether ungulates preferentially selected species containing lower phenolic levels, and also whether browsing induced either chemical or morphological changes in damaged plants. Phenolic levels varied greatly between plant species; ungulates browsed preferentially on the species with the lowest phenolic levels. No difference in phenolic content was found between browsed and unbrowsed trees. Morphological changes in heavily browsed trees included an increase in shoot and leaf density and a net decrease in leaf size. We suggest that for Mediterranean plants, which have evolved under high browsing pressure from large mammals, the production of small leaves and dense shoots in response to browsing might decrease ungulate foraging efficiency and hence reduce the rate of further damage as effectively as high levels of chemical defence.     

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.     

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.     

Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight

Abstract. We analysed the relationship between seed traits (weight, shape and dispersal structures) and the abundance and habitat segregation of Mediterranean grassland species. To take into account possible correlations with other plant traits, the study also includes 5 vegetative traits (growth form, plant longevity, clonality, onset of flowering and plant size) of commonly accepted functional importance. Data were recorded for 85 species from dehesa grasslands in central Spain. Species abundance was measured in upper (dry and less productive, high stress) and lower (moist and more productive, low stress) slope zones in the same area. Habitat segregation was estimated using an index based on the relative frequencies of species in upper and lower slope zones. Multiple regression models were fitted using species, as well as phylogenetically independent contrasts, as data points. Annual small‐seeded species without specialised dispersal structures are over‐represented in dehesa grasslands. Abundance was negatively related to seed weight in upper slope zones. None of the recorded plant traits were related to abundance in the lower slope zones. Habitat segregation was mainly related to seed weight, but also to some vegetative traits. Annual, early flowering and small‐seeded species were relatively more abundant in the upper than the lower slope zones. This pattern is independent of phylogeny. Our results suggest that in dry Mediterranean grasslands, abundance of many species is determined by dispersal (production of numerous small seeds) rather than by competitive ability.     

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.     

Fine-scale spatio-temporal patterns in a mountain grassland: do species replace each other in a regular fashion?

Abstract. Long-term ‘high spatial resolution’ permanent plot data were used to determine whether fine-scale species replacements in space occur more often than expected on the basis of random processes, and to test whether these replacements are species-specific. Monte Carlo tests were used. There was no indication of significance in associated positive change (species A increased when B increased); the overall number of significant results was not higher than expected on a random basis. For associated negative change, or replacements (species A increased when B decreased) the overall number of significant results was significantly higher than expected. Significant reciprocal replacements between Deschampsia flexuosa, Festuca rubra and Nardus stricta were frequent; changes in Anthoxanthum alpinum and Polygonum bistorta were uncorrelated with changes of the former three. The first three species thus use the ‘same’ space. Both the latter species often reproduce by seeds and their turnover is much higher. The prevalence of negative correlations of changes (i.e. correlation of increase with decrease) supports the concept of an internal structuring within the grassland community.     

Prediction of occurrence of vascular plants in deciduous forests of South Sweden by means of Ellenberg indicator values

Abstract. In this study we present a new method for predicting the occurrences of species using data from deciduous forests in South Sweden. Complete species lists of vascular plants were compiled from 101 stands and from representative sample plots inside the stands. Soil samples from each stand were collected for determination of pH and nitrogen mineralization. Presence-absence data for species were fitted to the values of four environmental variables - soil moisture, soil reaction (pH), soil nitrogen and light - by means of Linear (Multiple) Logistic Regression (LLR), and Gaussian (Multiple) Logistic Regression (GLR). First, these values were estimated by calculating the weighted averages of Ellenberg indicator values. Second, the estimates for reaction and nitrogen were substituted by the real measurements of pH and mineralized NH₄⁺, keeping the Ellenberg estimates for light and moisture. The models were validated by an independent test data set. In general, the models had high predictive abilities. GLR fitted the species occurrences better to the environmental variables than LLR, but had a lower accuracy of prediction of species occurrence in the stands. The use of soil measurements instead of Ellenberg indicator values did not improve the predictive abilities of the models. The environmental conditions in the stand test set were successfully estimated by using species data from the plots. When using the species lists of the stands instead of plot data, a slightly better predictive ability was obtained. The collection of plot data, however, is easier and less time-consuming. The accuracy of prediction differed considerably between species.